"STORY MUSEUf Bulletin of 26 NOV 1997 The Natural Histor Museum

Botany Series

THE NATURAL HISTORY MUSEUM

VOLUME 27 NUMBER 2 27 NOVEMBER 1 997 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum

(Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology.

The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Ms M.J. Short

Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance.

A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to:

Intercept Ltd. P.O. Box 7 16 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058

Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world.

World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.)

The Natural History Museum, 1997

Botany Series ISSN 0968-0446 Vol. 27, No. 2, pp. 77-147

The Natural History Museum Cromwell Road London SW7 5BD Issued 27 November 1997

Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. not. Hist. Mus. Loncl. (Bot.) 27(2): 77-147 Issued 27 November 1997

Systematics of Pogostemon (Labiatae) 78 G.R. BHATTI AND M. INGROUILLE

included three D. D. maintained as operational taxonomic units as a first step in a revision Dysophylla species: auricularia, myosuroides and D. Benth. In the same work Bentham of the infrageneric classification. Benth., strigosa (1832- divided the into two the Non-patchouli species of Pogostemon sensu stricto are more 1836) genus Pogostemon sections, basis of the of inflores- diverse and better delineated. For example there is P. litigiosus Doan Paniculatae and the Racemosae on the type with a two-lipped calyx. Species are generally narrowly endemic to cence. was taken areas such as Sri Lanka or southern India, the eastern Himalayas, Bentham's splitting of Dysophylla and Pogostemon Yunnan, Java, Borneo or southern Africa. further, first by Rafinesque-Schmaltz (1847) and later by Briquet raised Dysophylla includes all the small flowered species which tend to (1897). Rafinesque-Schmaltz (1847) effectively Dysophylla have flowers crowded into dense verticillasters. Many species have section Verticillatae to generic rank by publishing a new genus and M. a weak stem, probably because they are aquatic or marshland Eusteralis Raf, which included Mentha pumila Graham subdivided and inhabitants. The calyx is small, translucent, and infundibular with a verticillata Roxb. Briquet (1897) Dysophylla follows: wide diameter. Most species have verticillate, linear to linear- Pogostemon into sections and subsections, as lanceolate leaves. However, there are a number of species which Genus Dysophylla combine the character states of Pogostemon sensu stricto and Sect. l.Rhabdocalicinae Briq. (Calyx tube cylindrical and rounded Dysophylla, so that the boundary between these taxa has not always or very indistinct, pentagonal) been stable, with different workers putting the same species into one 1. Oppositifoliae Benth. or other group or not recognising the groups at all. This is described A. Perennial species in greater detail below. B. Annual species The study reported here is a re-examination of the variation of 2. Verticillatae Benth. Pogostemon sensu lato leading to the erection of a new infrageneric Sect. II. Goniocalicinae Briq. (Calyx tube prominently five- classification which is more natural and which may be used as the angled) basis for future studies at lower taxonomic rank. A cladistic and Genus Pogostemon phenetic analysis of characters has aided the resolution of new 1. Section Racemosa Benth. infrageneric taxa but this has not been used directly because certain A. Glabriuscula Briq. (Naked filaments) characters, thought to be more constant and reliable, have been B. Barbata Briq. (Hairy filaments) weighted as more important in the classification, but all characters 2. Section Paniculata Benth. were weighted equally in the cladistic and phenetic analysis. The A. Interrupted verticils cladistic and phenetic analysis will be reported elsewhere. B. Continuous verticils

History of the classification of Pogostemon Kudo (1927) treated Dysophylla and Pogostemon as distinct genera under subtribe Pogostemoninae. He divided Dysophylla into Pogostemon was described by Desfontaines (1815) as a distinct two sections: genus of Labiatae because of its hairy stamens: pogon is Greek for - beard and pogonostemon means 'with bearded stamen' (Steam, Section Eudysophylla stem indumentum tomentose to hirsute and 1992). The type species for the genus is P. plectranthoides Desf. leaves opposite, broad, ovate-lanceolate, margin serrulate (in- Desfontaines placed his new genus near Hyssopus L. Dysophylla cludes only D. auricularia). Blume was described as a distinct genus, related to Pogostemon by - Section Chotekia stem glabrous to pubescent and leaves verticil- having bearded stamens (Blume, 1 826). Dysophylla had been previ- late, narrow, linear to linear-lanceolate with an entire margin. ously recognized as distinct by some authors under different names. Hermann (1717) gave the polynomial Veronica hirsuta latifolia Four species which belong to Bentham's Dysophylla section Zeylanica aquatlca to a species of Dysophylla. Linnaeus (1747) Oppositifoliae (D. auricularia, D. myosuroides, D. rugosa Hook.f. listed a genus Alopecuro-veronica, a name he later cited as a andD. salicifolia Dalzell ex Hook.f.) were transferred toPogostemon synonym of Mentha auricularia (Linnaeus, 1767). The name on the basis of having opposite, broad and petiolate leaves, presence Majorana foetida was given as another synonym, a species de- of crystals in the calyx, and absence of an aerenchyma tissue in the scribed by Rumphius (1750) as Majana foetida. He described it as stem (El-Gazzar & Watson, 1967). Wu & Li (1975) also transferred having no close affinity to other species of Mentha. Blume ( 1 826) D. auricularia and D. falcata C.Y. Wu to Pogostemon but without cited Rumphius' illustration when he recognized Dysophylla as a mentioning any reason for doing so. The removal of D. auricularia, distinct genus based on Dysophylla auricularia (L.) Blume (= in particular, posed a nomenclatural problem because it is the type Mentha auricularia L.). Although he placed Dysophylla next to species ofDysophylla. A solution to this problem was put forward by Mentha L. he considered it different from Mentha by having closure Panigrahi (1976) and Bakhuizen van den Brink & van Steenis of the fruiting calyx, fleshy swelling of the disc, and bearded (1963), who suggested the generic name Eusteralis Raf. for the stamens. At about the same time Blume (1826) published a species remaining species of Dysophylla, those in section Verticillatae. from Java he called P. menthoides Blume. Later, however, Bentham Keng (1978) also placed Dysophylla section Verticillatae in (1829) questioned the placing of P. menthoides in the genus Pogostemon section Eusteralis Raf. However, Panigrahi (1984) Pogostemon because it had naked filaments. referring to Article 22.4 (I.C.B.N. 78) mentioned that Pogostemon Bentham (1829, 1830, 1832-1836) also widened the concept of section Verticillatae Benth. has priority over Pogostemon section Dysophylla by including species described as Mentha by Loureiro Eusteralis (Raf.) H. Keng. (1790) and Roxburgh (1814, 1832). Bentham (1830) divided Several authors found it difficult to distinguish Dysophylla from Dysophylla into two groups on the basis of phyllotaxy, one group Pogostemon (Hasskarl, 1842; Miquel, 1859; Kuntze, 1891; Keng, with opposite leaves and the other verticillate leaves. In Labiatarum 1978). Press (1982) criticized Briquet's division of Pogostemon and

et 1 genera species Bentham ( 832) formally recognized these groups Dysophylla because of the obscurity of the characters used. Press' as section and section Verticillatae. Section 1 for the Oppositifoliae ( 982) phenetic analysis of characters found no evidence Oppositifoliae, which agreed with Blume's original description of grouping of taxa as proposed in Briquet's classification. SYSTEMATICS OF POGOSTEMON 79

MATERIALS AND METHODS for 8-12 minutes. Almost all of the samples examined were air dried, but a comparison was made with fresh material, dried using a critical point drier (POLARON) to confirm characterization. All the listed in the account specimens systematic were examined Data were subjected to cladistic analysis and phenetic analysis and used for data analysis. Some other specimens were rejected (Bhatti, 1995). Details and a comparison of the results of different because they were incomplete at the correct state of development or cladistic and phenetic analysis will be reported elsewhere. had no information about their geographical origin. As a result the following possibly distinct species were not available: Pogostemon brevicorollus Y.Z. Sun, P. reticulatus Merr., P. falcatus (C.Y. Wu) RESULTS C.Y. Wu & H.W. Li, P. szemacensis (C.Y. Wu & S.J. Hsuan) Press, and P. tsiangii (Y.Z. Sun) Press. Herbarium material of Pogostemon For this most in floristic works and other genera has been obtained from the herbaria of The study species currently recognized were maintained. Three new are described below. Natural History Museum (BM), the Royal Botanic Gardens, Kew species Geographical coverage was patchy. In particular there were few (K), and the Royal Botanical Garden Edinburgh (E). More than half specimens from China and parts of southeastern Asia. Even in the of the total number of species under investigation have been re- Indian sub-continent, from where the of ported from India. A field excursion to this country was undertaken majority samples origi- nated, it was clear that some areas were much better in 1991. The following herbaria of the Botanical Survey of India geographical than others. This reflect in the floristic were visited: Poona (BSI), Calcutta (CAL), Coimbatore (MH), represented may part greater diversity of ofAssam and Keralia, but also reflects the Dhera Dune (BSD), and Rabinath Herbarium, St. Joseph College parts probably historical incidence of individual collectors. The lack of (RHT). sampling by a good sample of the variation of species made it at Wherever doubt existed regarding identification, type material many impossible this stage to consider redefining the limits of most species. Instead and the type description have been examined. In eighty-five per cent we have concentrated on describing reliable characters for identifi- of species it was possible to study at least one type specimen. A few cation in the systematic account below. Nonetheless it was clear that species have been observed in their natural habitat in India. These some species are more distinct than others. include: Pogostemon paniculatus (Willd.) Benth. (not flowering), P. An infrageneric classification is proposed which resurrects, at purpurascens Dalzell (not flowering), P. quadrifolius (Benth.) least in part, that of previous workers. However, the inclusion of new Kuntze, P. heyneanus Benth. (cultivated), P. auricularius (L.) Hassk., characters, especially those from a detailed study of floral morphol- and P. plectranthoid.es Desf. (not flowering). Cultivated material of ogy, has allowed subgenera, sections, and subsections to be defined P. plectranthoides and P. quadrifolius was available and used to more clearly. The order of taxa in the systematic treatment is similar check that dried material accurately represented the living state. to that in a computed consensus tree obtained by cladistic analysis Descriptions of species in the various Floras of the different but the arrangement is not identical because in the classsification the regions were also examined. However, it has been found that many characters have been weighted by their ease of use and reliability. floristic accounts simply repeat descriptions of earlier workers, even Complex character states and combinations of unitary character where these are erroneous. Some specimens have been regularly states which may be less likely to exhibit parallel or convergent misidentified because of the close similarity of a number of species. evolution have been used to define groups at the higher ranks in the For example, the names Pogostemon brachystachyus Benth., P. hierarchy and more simple unitary characters at lower ranks. There fraternus Miq., and P. macgregorii W.W. Sm. have often been is a shortage of 'good' characters and in some cases a polythetic wrongly applied by earlier workers. approach has been adopted as a more reliable guide to forming a Mature plant specimens were selected for examination. The natural group. In these cases rank has been established with refer- maximum size of each part of the plant were recorded. However, for ence to the magnitude of the taxonomic discontinuity between the many tall plants, the herbarium sheets have been prepared from only groups and the size of the group, validated where possible with the upper parts of the plant with the inflorescence. Much longer reference to geographical and ecological data. Some groups are leaves are present at the basal part of the plant. For example, in linked by intermediate species but these have been allocated to the Pogostemon plectranthoides older leaves towards the basal part of group which contains their closest relative. the stem of about 23 cm long have been observed in living material, whereas the maximum size in herbarium specimens is about 14 cm. of infrageneric classification The small size of the flowers posed many difficulties in carrying Summary out a detailed of the and corolla from herbarium study calyx directly Genus Pogostemon Desf. material. Flowers from 1.5-4 mm. Permanent slides of the ranged I. Subgenus Pogostemon sensu Bhatti & Ingr. corollas and and where bracts, were made in order calyces, possible II. Subgenus Allopogostemon Bhatti & Ingr., subgen. nov. to record some of the characters which have never unique previously a. Section Racemosus (Benth.) Bhatti & Ingr., stat. nov. been Dried obtained from herbarium fully described. flowers, speci- i. Subsection Racemosus were boiled in water. Then the soaked corollas and mens, calyces ii. Subsection Glabriusculus (Briq.) Bhatti & Ingr., stat. nov. were dissected to their inner sides and mounted on slides expose b. Section Zygocalyx Bhatti & Ingr., sect. nov. with DPX mountant without other treatment. any III. Subgenus Dysophyllus (Blume) Bhatti & Ingr., comb, et stat. Normally 5-10 calyces and corollas were examined from each nov. if there doubt or in the specimen, but, was any complication scoring a. Section Dysophyllus then the number of were increased. required information, samples b. Section Verticillatus (Benth.) Bhatti & Ingr., comb. nov. Nutlets were obtained from fruiting calyces. A scanning electron microscope (JEOL JSM 35CF) study was carried out on nutlet Names of infrageneric taxa used in the text below are sensu Bhatti & morphology and hairs at the middle and towards the lower part of the Ingr., except where stated otherwise. filaments. Samples were collected from herbarium sheets, mounted The shrubby patchouli species which have been placed here in include several rather defined taxa on stubs, and coated with gold by a sputter coater (EMITECH K550) subgenus Pogostemon poorly M. INGROUILLE 80 G.R. BHATTI AND

will be subdivided into several smaller taxa. Data to enable stable around Pogostemon benghalensis, P. plectranthoides, P. cablin, and to be erected are not available. P. heyneanus. Patterns of variation are complicated by the cultiva- groups yet tion of variants which are maintained vegetatively and rarely flower and their introduction into areas outside the native distribution of the New data are before be considered as species. required they may CHARACTER VARIATION variants of one or more polymorphic species. We have hesitated to submerge all species of subgenus Pogostemon into a single polymor- Indumentum and glands phic species P. plectranthoides sensu lato. based is is nature and distribution of the In subgenus Dysophylla a similar population study A significant feature of the genus the needed to evaluate the relationship of narrowly restricted endemics hairs. Most important are the moniliform hairs found on the fila- various other like Pogostemon erectus Kuntze, P. koehneanus (Muschl.) Press, ments of almost all species (Fig. 1). Hairs of kinds, taxa found and P. andersonii (Prain) Press to geographically widespread simple, stellate, fruticose, and dendromorphic, have been and P. kinds of hairs like P. stellatus (Lour.) Kuntze, P. crassicaulis (Benth.) Press, elsewhere on plants (Fig. 1). In most species similar the cruciatus (Benth.) Kuntze. can be found on all parts of the plant but rarely different parts of or branched. A newly defined group is Pogostemon subgenus Allopogostemon plant have different hairs. Hairs may be simple Many which consists of most non-patchouli species of Pogostemon sensu have a swollen basal cell which has different cell contents from the with well marked Section trichomes. Hairless stricto. It is a diverse group species. distal part. These may be glandular glands may five from Racemosus has eleven species in southern India, species be found intermixed with hairs (Fig. 1 ). of section the Philippines, and one from southern Africa. Species Simple trichomes are either unicellular or uniseriate multicellular. with P. Zygocalyx show a trend towards having a two-lipped calyx, They are elongated, broader at the base, and gradually tapering is centred on an area from litigiosus the most extreme. This section towards the apex. In multicellular types they are generally slightly but includes three the eastern Himalayas to Yunnan and Thailand swollen at the septa, forming a node. All taxa have multicellular, Sec- species endemic to Java, Borneo, and Sri Lanka respectively. simple trichomes except Pogostemon koehneanus and P. peguanus iionZygocalyx shares some characters with subsectionGlabriusculus (Prain) Press which have unicellular hairs on stems, leaves, and of section Racemosus. Pogostemon wightii Benth. of section inflorescence. Stellate trichomes have been found only in P. of subsec- Zygocalyx has a similar shaped corolla to that of species tuberculosus Benth. On most of the plant the hair branch which P. Benth. and P. tion Glabriusculus. The calyx of reflexus speciosus occupies the central position is 2-celled and longer than the others. Benth. of subsection Glabriusculus has some teeth rather subulate The remaining lateral branches, which are unicellular, radiate from and fringed by bristles like section Zygocalyx. The pattern of hairs the central one, at 30-40 to the surface. Stellate hairs on the calyx on the interior of the calyx of these two groups is homologous. have 2-3-celled central branches. Fruticose trichomes are composed Pogostemon paludosus Benth. of subsection Racemosus shares char- of a bunch of branches, attached only at the base. The central branch on acters with subsection Glabriusculus, by having stalked glands is longer than the lateral branches and together they look like a bush. a the nutlets, and section Zygocalyx, by having somewhat asym- This type of trichome is found in P. velatus Benth. and P. williamsii lacks metrical calyx, but it has moniliform hairs on the stamens and Elmer. Press (1982) reported these trichomes as sessile branched subulate teeth, the key characteristics of these taxa. hairs in comparison to dendromorphic trichomes because they are Most species of subgenus Dysophylla have a corolla in which the without a central boss and have long flexuous branches with each are separate lobes of the upper lip and the single lobe of the lower lip branch uniseriate and multicellular. However, the use of the term has a approximately equal, and are shaped so that the corolla sessile is unfortunate because this is impossible to determine with- linear-lanceo- symmetry which is nearly radial. Many have linear to out branching. Dendromorphic trichomes are branched hairs in useful characters but there are a number of late leaves. These are key which the basal part is like the main trunk of a tree. This basal part andersonii different species in section Verticillatus: Pogostemon gives rise to lateral branches of considerable length. The lateral (Prain) Press, which has two opposite leaves at each node and P. branches are arranged either in opposite or alternate fashion. Fruticose trinervis Chermsir. ex Press, which has orbicular leaves. and dendromorphic trichome types occur in combination in P. Section Dysophyllus is a poorly delimited group. It includes velatus and P. williamsii, although the leaves and calyces of P. species intermediate between section Verticillatus and subgenera williamsii have only dendromorphic trichomes. The abaxial leaf PogostemonandAllopogostemon. The boundary belweenDysophylla surface of P. elatispicatus Bhatti & Ingr., has both simple and sensu stricto has taxed workers. It has and Pogostemon previous only dendromorphic hairs, whereas other parts of the plant have only for a been resolved here by recognizing section Dysophyllus group simple hairs. of heterogeneous species which do not fit easily into either subgenus. Emphasis has been placed here on calyx characters so that species in Stem section Dysophyllus may have varied characters of Pogostemon sheets sensu stricto, but have the broadly conical and translucent calyx of Collectors' remarks on herbarium sometimes provide helpful stem is erect in most Dysophylla; so, for example, P. amaranthoides Benth. which has notes about the form and habit of the plant. The normally been placed within Pogostemon sensu stricto is here placed species. The exceptions are taxa like Pogostemon auricularius, in subgenus Dysophyllus. which has an aquatic habit. In nature the stems of this species are Pogostemon mutamba (Hiern) G. Taylor and P. micangensis G. prostrate and root at the nodes; it is a straggling plant and spreads with the erect Taylor, the African species, have also traditionally been placed in over an area of 1-2 m in marshy places flowering of the Pogostemon sensu stricto. They have the normal zygomorphic co- branches arising from the main stem in water. The part plant is branch but here rolla of other Pogostemon species from subgenera Pogostemon and which is used to prepare a herbarium sheet an erect Allopogostemon, but although they also have a calyx with intercostal it has been considered as an erect stem. P. rupestris Benth. roots at those veins it is very strongly conical/campanulate like species of subgenus the nodes. Many species of subgenus Dysophyllus, especially Dysophylla. of section Verticillatus, have a weak stem, sometimes contracted at In of Section Dysophylla is clearly not a natural group and in the future the internodes and obviously procumbent. contrast, species SYSTEMATICS OF POGOSTEMON 81

b

Fig. 1 Indumentum types observed in Pogostemon: (a) gland without a hair; (b-d) unicellular trichomes, (c, d) with basal glandular cell; (f-j) uniseriate

multicellular trichomes, (e) with basal glandular cell; (k) fruticose trichomes; (l-o) stellate trichomes, (1, n) with unicellular arms, (m, o) with some

multicellular arms, (1, m) sessile, (n, o) stalked; (p, q) dendromorphic, (p) branches opposite, (q) branches alternate; (r, s) moniliform filament hairs. 82 G.R. BHATTI AND M. INGROUILLE

Fig. 2 Leaf forms found in Pogostemon: (a) whorl of five leaves at a node; (b) whorl of three leaves at a node; (c) two leaves opposite at a node; (a, e, f)

linear to linear-lanceolate; (b, c, g-1) ovate to lanceolate; (d) pinnatifid; (1) base cordate; (b-k) base attenuate to truncate; (a, b, e-h) sessile to subsessile;

(c, h-1) petiolate. SYSTEMATICS OF POGOSTEMON 83

subgenus Pogostemon are shrubs or subshrubs. have a large semi-woody subsessile leaf and P. barbatus Bhatti & Ingr. and P. Most other are herbs. species amaranthoides are petiolate. Petiolate leaves are normal in other the of the stem towards the Normally upper part inflorescence subgenera with only P. nigrescens, P. strigosus (Benth.) Benth., and is more than lower region hairy parts. A study of hairs on the stem P. velatus having a subsessile leaf. showed a stem in glabrous Pogostemon glaber Benth., P.fraternus The leaves of Pogostemon nelsonii are glabrous but all other P. nelsonii Doan, P. travancoricus P. Miq., Beddome, pumilus species of subgenus Pogostemon are hairy. Most species of subgenus P. P. (Graham) Press, lythroides (Diels) Press, helferi (Hook.f.) Allopogostemon are hairy but P. travancoricus has glabrous leaves. P. P. Press, yatabeanus (Makino) Press, sampsonii (Hance) Press, P. Several species of subgenus Dysophyllus have glabrous leaves, stocksii P. linearis P. (Hook, f.) Press, (Benth.) Kuntze, pressii including P. pressii, P. helferi, P. pentagonus, P. sampsonii, and P. Panigrahi, and P. faurei (Leveille) Press. Stellate trichomes have Verticillatus. been found on P. tuberculosus. Fruticose and dendromorphic The abaxial leaf surface of many species is densely hairy when trichomes are found on P. velatus and P. williamsii. The remaining compared to the adaxial surface. The veins are more hairy than the taxa have simple hairs. Unicellular hairs have been found on the lamina on both the adaxial and abaxial surfaces. stems of P. peguanus and P. koehneanus. Inflorescence Leaves The inflorescence in Pogostemon, like most Labiatae, is a terminal, verticillate This has a Inflorescences Most species have a pair of opposite leaves at each node but among thyrse. spike-like appearance. have been into three kinds: a terminal a the species of section Verticillatus the number of leaves at each node categorized single spike, terminal with two lateral and a terminal with ranges from three to more than ten, with four as the most common spike spikes, spike more than two lateral spikes 3). number. The number of leaves is normally constant within species (Fig. In the inflorescence is branched; the termi- but variation has been noticed in Pogostemon quadrifolius which has subgenus Pogostemon nal verticillaster is more than two lateral most nodes with whorls of four leaves, but two leaves at some nodes. usually accompanied by verticillasters, less two but in P. Pogostemon yatabeanus, P. sampsonii, P. crassicaulis, P. koehneanus, frequently by just laterals, it is Branched inflorescences are found in a and P. trinervis have three leaves in a whorl. Pogostemon cruciatus purpurascens simple. few in other but most have a verticillaster. and P. stocksii have five leaves in a whorl. Eight or more leaves are species subgenera, single The verticillaster is dense but in some the present in P. erectus, P. deccanensis (Panigrahi) Press, and P. commonly very species verticils stellatus. lowest are arranged laxly and the upper part densely. A few species, in have a lax All species have simple leaves, except Pogostemon tisserantii especially subgenus Allopogostemon, verticillaster throughout. (Pellegr.) Bhatti & Ingr., which has pinnatifid leaves. Leaves may be ovate, lanceolate, orbicular, cordate, elliptic, ovate-lanceolate, or Bracts and bracteoles linear (Fig. 2). The majority of the members of subgenera Pogostemon and Allopogostemon have an ovate to lanceolate leaf shape, although The arrangement, shape, and persistence of the bracts and bracteoles P. rotundatus Benth. has orbicular leaves and P. speciosus has is very variable, even on a single plant. In some species bracts are large cordate leaves. Species to belonging subgenus Dysophyllus usually and there is a continuum in size and shape between lower vegetative have linear to linear-lanceolate leaves. Most are found in exceptions leaves on an inflorescence and the bracts associated with the verticils. sectionDysophyllus: P. auricularius, P. (Benth.) Kuntze, myosuroides This pattern is common in subgenus Pogostemon. More frequently and P. salicifolius Dalzell ex Hook.f.) El Gazzar & L. Watson have bracts are distinctly different from vegetative leaves: smaller, nar- a lanceolate to linear-lanceolate leaf while an leaf shape elliptic rower, and less petiolate. Sometimes they are indistinctly toothed. In 3 has been recorded from P. mutamba and/ . shape glabratusChermsir. P. purpurascens, P. cablin, P. nepetoides, and P. nelsonii they are ex Press. trinervis is unusual in section Verticillatus Pogostemon distinctly toothed. Deeply cleft, two-lobed bracts can be found in P. because its leaves are more or less orbicular. Most have a species villosus Benth. andP.paniculatus. In subgenus Pogostemon the bracts cuneate leaf base but P. has leaves which are cordate at the speciosus and bracteoles are smallest and narrowest in P. tuberculosus, P. base. Leaves with a rounded base have been found inP atropurpureus formosanus, P. elsholtzioides Benth., and P. dielsianus Dunn. Bracts Benth., P. P. P. mollis Benth., P. brachystachyus, macgregorii, in the other subgenera are normally narrow, linear-lanceolate to linear, philippinensis S. Moore, P. pressii, and P. trinervis. and filiform bracteoles are found. In section Verticillatus several The leaf is acute or acuminate, but an obtuse apex commonly apex species, including P. helferi, P. pumilus, P. stocksii, P. erectus, P. has been recorded from P. cablin, P. Pogostemon brachystachyus, stellatus, and P. deccanensis the bracts/bracteoles are spathulate. P. mollis, P. P. Dunn, P. glaber, nepetoides Stapf., nigrescens Bracts and bracteoles are normally densely hairy. rotundatus, P. speciosus, P. velatus, P. peguanus, P. pentagonus (C.B. Clarke ex Hook.f.) Kuntze, P. quadrifolius, P. sampsonii, P. Calyx trinervis, and P. cruciatus. Dentate, double dentate, crenate, double crenate, inciso-crenate, The calyx is either infundibular, tubular or campanulate. In subgenus and serrate margins have been recorded. An entire/revolute margin is Dysophyllus the calyx is infundibular with a wide diameter, greater found in some species of subgenus Dysophyllus, but not elsewhere. than the length of the tube. In subgenus Allopogostemon the calyx is The leaf margin in Pogostemon cristatus Hassk., P. formosanus normally tubular or infundibular, and then two-lipped and with a Oliver, and P. gardneri Hook.f. has a deep incision, and is described circumference less than or equal to the length of the calyx tube. In as inciso-crenate. subgenus Pogostemon the calyx is normally infundibular and has a Petiolate, subsessile (petiole < 5 mm), and sessile leaves have diameter narrower than the tube length. been observed. Most of the members of subgenus Dysophyllus have Pogostemon normally has a calyx with five teeth. Some speci- a sessile leaf, a character state not found outside this subgenus, mens ofP. atropurpureus, P. brachystachyus, P. micangensis, and P. although Pogostemon auricularius, P. glabratus, P. mutamba, P. philippinensis have 5-7 toothed calyces. Extra teeth are particularly myosuroides, P. micangensis, P. quadrifolius, and P. salicifolius common in P micangensis so that the calyx toothing is rather 84 G.R. BHATTI AND M. INGROUILLE

3 Inflorescence in Fig. types observed Pogostemon; (a, d, g) single spike; (b, e) with two lateral spikes; (c, with branched lateral spikes; (b, c, f, g) spikes dense; (a) spike lax below, dense above; (d, e) lax. SYSTEMATICS OF POGOSTEMON 85

some of the have four teeth. irregular. InP.faurei calyces Calyx teeth more or less distally, in some species, just to the tip of the teeth, as either or are normally broadly narrowly triangular. in Pogostemon membranaceus Merr. and P. petiolaris Benth. of section Species Zygocalyx have the most distinct calyx. The An alternate pattern of hair distribution, which can be confused calyx is asymmetrical and two-lipped, with the two lower teeth with the last condition, is where the hairs form a wavy annulus around narrower than the upper three, and the teeth tips drawn out into a the top ofthe calyx tube. In these species the most dense hairs are often This narrow point (subulate). condition is most pronounced in found at the teeth sinuses.The annulus varies from being very strongly but is seen to a lesser in Pogostemon litigiosus extent P.fraternus, P. developed mPogostemon speciosus, so that tufts of hairs seem to spill brachystachyus, P. wightii, P. nigrescens, P. strigosus, P. menthoides, out at the teeth sinuses, to poorly developed in P. strigosus, which has P. hirsutus Benth., and P. macgregorii. only a few hairs present, mainly at the sinuses of the teeth. In section Calyx venation is a very important character for delimiting Zygocalyx an annulus of hairs has been observed in the throat of the subgeneric groups. All species have five prominent veins along the calyx in P. wightii, P. brachystachyus, P.fraternus, P. litigiosus, P. ribs of the calyx. In subgenus Dysophylla the only veins which are macgregorii, P. menthoides, and P. nigrescens. This character state obvious are those at the ribs of the calyx and the intercostal region is may be homologous to that seen in section Racemosus subsection thin and translucent. Pogostemon micangensis has the same pattern Glabriusculus, in P. atropurpureus, P. reflexus, and P. speciosus but with irregular toothing so that there may be an intervening rib which have tufts of hairs at the tips of the teeth and a line of hairs which does not end in a tooth. In subgenus Pogostemon, as well as running down the edge of the teeth to the sinus. the five main rib veins, there are numerous secondary veins or lines The patterns described above are clearly different from that seen of elongated cells in the intercostal region, running in parallel to the in subgenus Pogostemon, where hairs are more evenly distributed ribs, and converging at the teeth sinuses. In the distal part of the tube within the calyx, over the teeth, and sometimes down into the tube and in the teeth secondary branches arise from the rib vein and run including the region of the ribs. Normally the upper part of the calyx to the thickened margin of the calyx. The calyx tube is incrassate and tube and teeth are hairy or only the teeth are hairy. Pogostemon opaque with the exterior often densely covered with hairs. In subgenus paniculatus, P. pubescens Benth., P. purpurascens, and P. wattii Allopogostemon two patterns may be observed: either the calyx has C.B. Clarke have the entire calyx tube and teeth hairy. Pogostemon five midrib veins plus five other strong intercostal veins, terminating gardneri has the tube but not the teeth hairy. at each sinus; or there is a more reticulate pattern of venation with In subgenus Dysophyllus the interior of the calyx is glabrous five rib veins plus some other intercostal veins. In the intercostal except for a few species which have the inner teeth hairy. region the number of veins is variable and the veins interconnect in Corolla a reticulate fashion with the rib veins. The intercostal region is thinner than in but rather than trans- subgenus Pogostemon, papery The corolla consists of two parts, the tube and the two-lipped limb. lucent as in subgenus Dysophyllus. The upper lip is three lobed. The outer surface of the upper lip, and hairiness is an character. Calyx important Pogostemon sometimes also the lower lip, is normally hairy. Pogostemon elsholtzioides, P. nelsonii, P. nepetoides, and P. travancoricus have tuberculosus, P. auricularius, P. rogersii N.E. Br., P. mutamba, and to the teeth. a of hairs on the a glabrous margin The presence of row P. paludosus have hairs only on the central lobe of the upper lip. inner side of the margin has been recorded in P. dielsianus and P. The corolla tube varies in length from shorter to longer than the formosanus. Hairs originating slightly inside and outside the margin limb. The tube may have a narrow diameter relative to its length and of the teeth have been recorded inf. philippinensis, P. strigosus, and be uniformly tubular or become wider in its distal part. The relative similar to the P. tuberculosus. Hairs on the margin are either those on size of the lower and upper lips and the size and shape of the upper exterior of the calyx and ciliate, or they are stiff and bristle-like. The lobes provide useful characters for distinguishing species. The

latter are found in section Zygocalyx. dissected corolla reveals that there are several prominent veins ( 1 2 to The exterior of the calyx is normally hairy to very hairy. In most 16) in the tube. Four of these give rise to staminal filaments. The species it is the distal part of the exterior calyx tube which is hairy. remaining veins continue as midribs or lateral veins of the lobes in P. ciliatus Pogostemon auricularius, P. barbatus, P. glabratus, and the upper and lower lip of the corolla. Bhatti & Ingr. have just the teeth hairy andPogostemon travancoricus The site of filament attachment in the corolla tube is closely is glabrous. correlated to the total length of the corolla. For example, the total The majority of the hairy species have simple, uniseriate, multi- length of the corolla in Pogostemon amaranthoides is about 2.6 mm cellular hairs in which the basal cells are broad and apical cells and the tube length up to the site of filament attachment is about 1.0 sharply pointed. Both fruticose and dendromorphic trichomes have mm. In P. velatus corolla length is about 8.0 mm and the site of been found in Pogostemon velatus, while in P. williamsii, which is filament attachment is 3.5 mm up the tube. The length of the exserted considered by various authors as synonymous with P. velatus, only part of the filament is more or less constant within species but varies fruticose trichomes like those on its leaves are found. Pogostemon between species, ranging from hardly exserted up to 5.0 mm. tuberculosus possesses stellate trichomes. These trichomes differ The presence of hairy filaments gave Pogostemon its name. hairs are from those on other parts of the plant, having a central 3-celled, Moniliform hairs are found on all four filaments. The rather than 2-celled, branch. In subgenus Dysophyllus most of the unilaterally distributed. Moniliform hairs are only absent from taxa have simple, multicellular hairs except P.faurei, P. koehneanus, section Glabriusculus, namely P. atropurpureus, P. speciosus, P. and P. peguanus, which have unicellular hairs on the outer surface of travancoricus, and P. reflexus, but these have a few simple hairs with the calyx. In subgenus Dysophyllus hairless glands are obvious as scattered over the filaments. Simple hairs are found mixed has a little patches of circular cells with yellowish cell contents. These are moniliform hairs in P. hirsutus. Pogostemon menthoides only probably present on all species but are not so obvious on the thicker few moniliform hairs present. filaments are nor- and opaque calyces of subgenera Pogostemon and Allopogostemon. In subgenera Dysophyllus and Pogostemon Observation of hairs within the calyx is difficult because they can mally glabrous in the lower part and at the base but a few simple hairs have be very fine and sparsely distributed here. Where visible, the distri- may be found here. Most species in subgenus Allopogostemon villous short hairs. The bution of hairs in the distal part provides an important character. The the filament base with sharp central/upper than the outer/lower two. hairs may be spread over the teeth and down the tube or confined two filaments are generally more hairy M. INGROUILLE 86 G.R. BHATTI AND

Gynoecium Nutlets under examination is terminated a bifid characteris- The style in all taxa by Only minor variations were observed in nutlet surface lobes. uniseriate 2-3-celled nutlets from stigma with equal or unequal Simple, tics within species (Figs 14-40). The majority of on the lobes of ciliatus. to hairs have been observed stigma Pogostemon species belonging to subgenus Dysophyllus are oblong. Ellipsoid of lobes is 1 .9-2. 1 recorded from P. linearis and The maximum length mm, ellipsoid-oblong nutlets have been found \nPogostemon flowers with both nepetoides. On one specimen of P. elsholtzioides P. verticillatus. Orbicular and ovoid nutlets have been found in P. have been observed. The flowers with 5 bifid and three-lobed stigmas deccanensis and/ , tisserantii respectively. In subgenera Pogostemon three-lobed stigmas have six nutlets. and Allopogostemon a wide range of shapes has been observed: Some glands have been found on the ovary in Pogostemon ovoid, orbicular, oblong, obovoid, lanceolate to linear-lanceolate, P. P. and P. tmvancoricus. and atrvpurpureus, reflexus, speciosus, ellipsoid to ellipsoid-oblong, ellipsoid-fusiform, 'D'-shaped. of the fit. The disc is four-chambered where each ovary segments A scanning electron microscope study of the nutlet surface re- in in all taxa. These chambers are normally equal and regular shape vealed several different types of nutlet epidermal structure (Table 1). with one arm of the disc has been surface is An asymmetrical disc, elongated, Few patterns correlate closely with groups. A smooth and P. stellatus of found in Pogostemon erectus, P. deccanensis, common in species of subgenus Dysophyllus. Subsection subgenus Dysophylla. Glabriusculus is characterized by having glands on the nutlets, a character also found in Pogostemon paludosus. Table 1 Nutlet surface morphology in Pogostemon. Nutlet colour could not always be ascertained because very mature specimens are required and of course mature nutlets fall out Nutlet surface morphology Species of the flowers.

smooth P. helferi, P. erectum, P. salicifolius, P. stellatus, P. pressii, P. peguanus, P. trinervis, P. stocksii SYSTEMATIC ACCOUNT P. P. puncticulate P. amaranthoides, benghalensis, (smooth surface with tiny gardneri, P. glaber, P. hispidus, P.

. nat. Paris scattered dots) paniculatus, P. plectranthoides, P. POGOSTEMON Desf in Mem. Mus. Hist. pubescens, P. villosus 2: 154(1815). smooth-lineate P. koehneanus Inflorescence a single, spike-like verticillaster or with two or more (thin lines running in parallel on lateral verticillasters. Leaves sessile or Bracts and bracteoles the smooth surface) petiolate. not membranous, 5-toothed; veined within, glabrous smooth-reticulate P. deccanensis persistent. Calyx or or outside. Corolla 4-lobed, (smooth surface is covered by a hairy within, densely sparsely hairy exterior of thin lined network) three lobes forming upper lip, lower lip single-lobed; upper or undulate P. crassicaulis lip densely or sparsely hairy, lower lip glabrous sparsely hairy. at different Filaments in two pairs, inserted at the same height or foveate (pitted) P. hispidus anthers heights in the tube, mostly unequal in length, exserted; smooth with thinly scattered P. nelsonii hairs towards the middle unilocular ; moniliform present (most species) small pits or with a few fine simple hairs, basal part glabrous or with a few simple reticulate with punctate walls P. purpurascens hairs, or densely villous, and then all bases equally hairy or only two P. pumilus, P.faurei reticulate-punctate lobes. Disc hairy. Style exserted; stigma with two equal or unequal granulate P. tuberculosus, P. verticillatus symmetrical or asymmetrical. Nutlets 4, rarely one. (evenly covered with closely spaced grains but without reticulate appearance) Key to subgenera, sections, and subsections reticulate (unornamented) P. auricularius, P. brachystachyus, P. than < 4 five 1 . Calyx broadly conical, diameter greater length, length mm, membranaceus, P. nigrescens, P. veins thinand translucent and/? mutamba); P. hirsutus only, (except/? micangensis philippinensis, < calyx teeth normally 1 mm; leaves normally sessile or subsessile reticulate with P. P. P. secondary nilagiricus, petiolaris, strigosus, 2 (petiole < 5 mm) (III. subgenus Dysophyllus) reticulations P. wightii diameter narrower than > 4 mm, five main reticulate-foveate with warty P. cristatus Calyx tubular, length, length veins but with intercostal thick or teeth patches veins, papery; calyx normally 3 > 1 mm; leaves normally petiolate rugose P. aquaticus base rounded or zig-zag channels P. ciliatus 2. Leaves linear or linear-lanceolate, sessile, truncate, cuneate, verticillate, > 2 per node (rarely 2 opposite leaves at each node) ruminate, penetrated by irregular P. delsianus, P. elsholtzioides, P. Illb. section Verticillatus channels running in different griffithii directions, showing eroded base Leaves lanceolate, elliptic, ovate, or clavate, petiolate, usually features cuneate, rarely rounded, 2 opposite leaves at each node (rarely some covered more or less P. rotundatus smooth, by nodes with 4) Ilia, section Dysophyllus circular scales 3. Calyx with 5 main rib- veins and multiple parallel intercostal secondary small tubercles P. myosuroides, P. elatispicatus, veins meeting at the sinuses; calyx tube often with scattered hairs within; P. strigosus, P. velatus, P. williamsii, bracts and bracteoles large, equalling or exceeding calyx, frequently P. quadrifolius broadly lanceolate or bifid and toothed or lobed, less frequently nar- stalked, cup-like glands P. speciosus, P. reflexus,P. rowly lanceolate; lateral verticillasters 2 or more, or rarely anunbranched travancoricus, P. atropurpureus, P. verticillaster but then filament bases normally glabrous or with long paludosus simple hairs I. subgenus Pogostemon SYSTEMATICS OF POGOSTEMON 87

with 5 main rib veins isolated, more or less branched 13. Calyx plus Calyx strongly asymmetrical and 2-lipped; filament bases glabrous ... intercostal veins from sinuses of teeth; annulus of hairs around present 41.P.litigiosus top of calyx tube and teeth; bracts and bracteoles linear to linear- Calyx weakly asymmetrical and filament bases villous lanceolate, large or small; inflorescence an unbranched verticillaster, or 2-lipped; 44. P.wightii if branched then filament bases glabrous or densely villous(II. subgenus Allopogostemon) 4 14. Calyx infundibular, with 5 main veins only, translucent 15

4. Calyx more or less 2-lipped; at least 2 teeth the calyx awl-shaped, Calyx with 5 main veins and intermediate veins running to the sinuses, margins fringed by stiff bristle-like hairs lib. section Zygocalyx texture thick 20

Calyx symmetrical; calyx teeth triangular with a fringe of fine hairs (Ha. 15. Corolla lower lip shorter than upper; filaments arising at two levels ... section Racemosus) 5 55. P. micangensis

5. Moniliform staminal hairs present Ilai. subsection Racemosus Corolla lips equal in length; filaments arising at the same level 16

Moniliform staminal hairs absent Haii. subsection Glabriusculus 16. Petiole > 5 mm long 50. P. barbatus

Petiole < 5 mm 17 Key to species long

17. Margin of calyx teeth glabrous 49. P. glabratus 1. Leaves in opposite pairs 2 Margin of calyx teeth fringed with white hairs 18 Leaves in whorls of three or more, at least at some nodes 59 18. Inflorescence < 30 mm long 62. P. andersonii 2. Inflorescence an unbranched verticillaster 3 Inflorescence > 30 mm long 19 Inflorescence branched, with two or more lateral verticillasters 32 1 9. Exterior of calyx very hairy with dense long hairs at base, teeth hairy 3. Stamens lacking moniliform hairs; ovary/nutlets with cup-shaped glands within 53. P. salicifolius 4 Exterior of calyx with few hairs but many glands, teeth glabrous within Stamens with moniliform and nutlets hairs; ovary usually without cup- 48. P. auricularius shaped glands 7 < 20. Calyx 4 mm long; length of corolla tube less than or equal to 4. Inflorescence lax; calyx exterior glabrous 36. P. travancoricus circumference 21

Inflorescence dense; exterior 5 calyx hairy Calyx > 4 mm long; length of corolla tube greater than circumference 23 5. Petiole > 4 cm long; hairs on stem and inflorescence 8-9 celled 35. P. speciosus 21. Inflorescence > 50 mm long 34. P. paludosus

Petiole < 4 cm hairs on stem and inflorescence 5-6 celled 6 long; Inflorescence < 50 mm long 22

6. An annulus of hairs in the teeth calyx throat, calyx conspicuously hairy; 22. Stem hairy; inflorescence lax below, dense above 23. P. mollis filaments inserted at the same height 37. P. reflexus Stem glabrous except at the nodes; inflorescence lax throughout An annulus of hairs around the tufted at calyx throat, conspicuously 26. P. rupestris teeth sinuses; filaments inserted at different heights 23. with 5 rib veins and minor veins 38. P. atropurpureus Calyx parallel secondary 8. P. purpurascens 7. Calyx tubular-inflated, asymmetrical, with 5 main veins and a variable with 5 rib veins and 5 sinus veins 24 number of intermediate, reticulately-branched sinus veins; teeth un- Calyx equal, lanceolate-subulate, with a margin of stiff bristles 8 24. Calyx teeth shallowly triangular 25

Calyx infundibular, tubular or tubular-inflated but symmetrical, with 5 Calyx teeth deeply triangular 29 main rib veins, with or without sinus veins; teeth equal, with or without of and fruticose nutlet surface a margin of fine white hairs 14 25. Indumentum dendromorphic hairs; spinulose 26 8. Inflorescence dense, continuous 9 Indumentum of simple, multicellular hairs; nutlet surface reticulate to with at least the lower verticil from the others Inflorescence separate reticulate-foveate 28 11 26. Petiole c. 5 mm long 30. P. velatus 9. Leaf margin dentate; calyx < 4 mm long; corolla < 5 mm long; moniliform hairs few on filaments 40. P. menthoides Petiole > 10 mm long 27

27. Inflorescence > 150 mm verticils and Leaf margin serrate; calyx > 4 mm; corolla > 5 mm long; moniliform long, arranged evenly relatively hairs dense on filaments 10 laxly throughout spike 33. P. elatispicatus

Inflorescence < 150 mm lax below or dense 10. Leaves subsessile, lanceolate; petiole < 5 mm long; inflorescence > 50 long, throughout 31.P.williamsii mm long 47. P. strigosus

> < 28. Inflorescence > 100 mm lax below, dense above Leaves petiolate, ovate; petiole 5 mm long; inflorescence 50 mm long, 29. P. long 43. P. hirsutus philippinensis

Inflorescence < 100 mm dense 24. P. vestitus 11. Inflorescence > 100 mm long 45. P. brachstachyus long, throughout

29. Leaves orbicular 28. P. rotundatus Inflorescence< 100 mm long 12 Leaves ovate 30 12. Verticils arranged densely in upper part 42. P. macgregorii

30. Corolla > 7.5 tube and narrow, c. 2 x limb Verticils arranged laxly throughout 13 mm long; long M. INGROUILLE 88 G.R. BHATTI AND 49 32. P. membranaceus 48. Lateral Verticillasters two

52 x limb .. 3 1 Lateral Verticillasters three or more Corolla < 7 mm long; tube relatively short and broad, < 2

49. Stem and leaf hairs < 500 um 50 31. Calyx c. 4.5 x 3.5 mm, with 10 main veins 22. P. nilagiricus 3-celled, long

indistinct Stem and leaf hairs 5-6 celled, > 500 5 1 Calyx c. 5.0 x 5.2 mm, with 5 main veins, subsidiary veins um long 25. P. petiolaris exterior 50. Stem dark reddish purple; petiole up to 40 mm long; calyx 33 13. P. glaber 32. Leaves subsessile, petiole < 5 mm long sparsely hairy

> 36 Stem to 10 mm exterior strongly Leaves petiolate, petiole 5 mm long green; petiole up long; calyx 20. P. hairy griffithii 33. Verticillasters lax below, dense above 46. P. nigrescens 51. Stem reddish; stem and leaf hairs > 1000 \im long 6. P. gardneri Verticillasters dense throughout 34 Stem green; stem and leaf hairs < 1000 pm long 14. P. hispidus 34. Verticillasters > 70 mm long 54. P. quadrifolius 53 52. Calyx glabrous within Verticillasters < 70 mm long 35 54 Calyx teeth hairy within 35. Calyx < 2 mm long; corolla < 3 mm long 52. P. myosuroides 53. Stem and leaves glabrous 1 2. P. nelsonii > P. Calyx > 2 mm long; corolla 3 mm long 56. mutamba Stem and leaves with stellate hairs 19. P. tuberculosus 36. Inflorescence lax, at least below 37 < the 54. Calyx teeth shallow; lower lip of corolla relatively narrow, upper Inflorescence dense 48 55 lip

37. Inflorescence with two lateral Verticillasters only > the Calyx teeth deep; lower lip of corolla relatively wide, upper lip 51. P. amaranthoides 56

Inflorescence with more than two lateral Verticillasters 38 55. Corolla up to 8 mm long, lower lip > 2 mm long

. 3. P. plectranthoides 38. Calyx 10-veined, lobes unequal 39

.... 4. P. cells in Corolla up to 6.5 mm long, lower lip < 2 mm long pubescens Calyx with 5 main veins and numerous parallel veins or lines of intercostal region, lobes equal 40 56. Filaments arising at two heights 57 of hairs in the throat 39. Plant upright, up to 1 m tall; calyx with an annulus Filaments arising at the same height 58

39. P. fraternus 57. Calyx tube sparsely hairy within 1 1 . P. heyneanus Plant procumbent, rooting at nodes; calyx lacking an annulus of hairs in Calyx tube glabrous within 21. P. dielsianus the throat 27. P. rogersii 58. Calyx c. 3 mm long; corolla < 4 mm long 15. P. wattii 40. Indumentum on stem and leaves with 2-3-celled hairs

17. P. formosanus Calyx c. 4 mm long; corolla > 5 mm long 2. P. villosus

Indumentum on stem and leaves with more than 3 cells 41 59. Leaves linear to linear-lanceolate 60

41. Petiole c. 6 mm long 18. P. elsholtzioides Leaves lanceolate 68. P. koehneanus

> 10 42 Petiole mm long Leaves pinnatifid 57. P. tisserantii

42. Inflorescence normally < 50 mm long 43 Leaves orbicular 72. P. trinervis

44 Inflorescence normally > 50 mm long 60. Leaves > 6 at each node 61

16. P. 43. Lateral Verticillasters with a single verticil champion!! Leaves < 6 at each node 64

Lateral Verticillasters with several verticils 45 61. Disc regular 75. P. stocksii

44. Stem weak, contracted at internodes; corolla up to 1 mm long Disc irregular, with one arm elongated 62 5. P. cristatus 62. Calyx teeth hairy within 76. P. erectus Stem strong and even; corolla < 10 mm long 46 Calyx glabrous within 63 45. Verticils surrounded by broad ovoid bracts, some deeply cleft of filament 7. P. paniculatus 63. Corolla very short, < 2 mm long, exserted portion exceeding it in length 77. P. stellatus Verticils surrounded by elliptic-oblong bracts, sometimes toothed but

of filament it in . never deeply cleft 9. P. cablin Corolla > 2 mm long, exserted part equalling length 78. P. deccanensis 46. Corolla < 5 mm long; nutlet surface foveate 14. P. hispidus

.. 65 64. Corolla with middle lobe of upper lip narrower than lateral lobes Corolla > 5 mm long; nutlet surface puncticulate 47 Corolla with middle lobe of upper lip wider than lateral lobes 66 47. Corolla 8-9 mm long; leaves with margin double dentate, apex acumi- < 61. P. nate, base cuneate; hairs on all parts of plant < 500 um long 65. Inflorescence 40 mm long pentagonus 1. P. benghalensis Inflorescence > 40 mm long 67. P. peguanus Corolla 6-7 mm long; leaves with margin inciso-serrate, apex obtuse, 66. Stigma hairy 59. P. ciliatus base truncate;hairs on all parts of plant > 500 um long 10. P. nepetoides Stigma glabrous 67 SYSTEMATICS OF POGOSTEMON 39

67. Inflorescence with more than two lateral verticillasters 68 more than two lateral spikes; hairs 5-celled, 1 100 um long; bracts ~ ~ ovate ' 6 7 x 2 3 mm ' hairv - Cal x tubular-inflated, 4.3 x 4.0 5- Inflorescence unbranched or with 2 lateral verticillasters only 69 y mm, veined; teeth and of tube outside, within; < upper part hairy glabrous olla 2 mm long 70. P. verticillatus teeth dliate; longest tooth c j 2-l .3 mm long, 0.8-0.9 mm wide at bas outer Corolla > 2 mm long 79. P. aquaticus e; hairs 2-celled, c. 200 um long. Corolla up to 8.7 mm lower c. 2.2 x 2 c. , long; lip mm; upper lip 2 mm across; central lobe 69. Inflorescence with two lateral verticillasters ... 69. P. crassicaulis only , m c. 1 .6 x 0.7 mm; lobes hairy outside and scattered down the tube. Inflorescence unbranched simple, 70 Filaments inserted at different heights, the lowest at a height of 4 mm

*n the tube; fi' aments c- 5.1-7.0 mm exserted c. 2.3 70. Leaves in whorls of 3 71 long, portion mm; two filaments glabrous towards the base. Style c. 9.5 mm long; Leaves in whorls of 4-5 stigma lobes c. 1.2mm.Discc. 1 mm long. Nutlets 4, c. 1200x700

m > 7 1 . < obovoid, Stem sparsely hairy; inflorescence 35 mm long .... 64. P. sampsonii M puncticulate. DISTRIBUTION. Stem glabrous; inflorescence > 35 mm long 65. P. yatabeanus Pogostemon benghalensis grows in open Bombax- Trewia riverine forest on the bare flood plain in Nepal and is also 72. lower corolla than , , . Calyx asymmetrical; lips larger upper 73 f , D , , D T ., i found in India, Bangladesh, Burma, Thailand, Sri Lanka, and China.

Calyx symmetrical; lower corolla lips equal to or less than upper .. 74 ETHNOBOTANY. One of the patchouli species. Pogostemon 73. Corolla < 1 mm long 71.P.pumilus benghalensis contains an astringent resin, an alkaloid, a yellow varnish of a slightly bitter taste and mouse-like odour of Corolla > 1 mm long 73 P. helferi trimethyl- amine, and a volatile oil with an odour like that of cedar wood. 74. Leaf base rounded or truncate 75 Extracts are a stimulant and styptic. Fresh leaves are used to clean Leaf base cuneate 76 wounds and provide healthy granulation. P. benghalensis is a good source of or nectar for < pollen bees, thereby providing panagol honey. 75. Inflorescence 20 mm long 58. P. lythroides SPECIMENS EXAMINED. Kerr 10333 (BM), Kerr 17496 (BM), Inflorescence > 20 mm long 74. P. press,, Kingdon . Ward 20299 (BM K)> Tmfh 67? (BM)> ^ 75Q (BMX 76. Leaf margin entire, revolute 63. P. cruciatus Wang 30222 (BM).

Leaf dentate 77 margin Pogostemon benghalensis has previously been recorded as a syno- n m of R but are distinct. However, it is 77. Stem rooting at nodes, reddish brown; leaves sparsely hairy y plectranthoides, they 60. P. linearis possible that many records of this species from southern India and Sri Lanka are referable to P. plectranthoides. Pogostemon . . Stem not rooting at nodes, not reddish brown; leaves glabrous , , , , ,, ,, ,. , benghalensis has a corolla with a narrower cylindrical tube and a 66. P. taure, ...... verticillaster which is generally less crowded than in P. plectranthoides It is a variable species with smaller flowered vari- I. Subgenus POGOSTEMON sensu Bhatti & Ingr. ants (= P. parviflorus Benth.) with verticillasters less than 10 mm in Pogostemon subgenus Paniculatae Benth., pro parte. diameter and purple-stemmed plants (= P. purpuricaulis Dalzell).

Stem strong; leaves in opposite pairs at each node, distinctly peti- 2. Pogostemon villosus Benth., Labiat. gen. spec.: 153 (1833). olate (P. griffithii sessile), ovate, rarely lanceolate; inflorescence a Figs4b, 14b. verticillaster with two or more lateral verticillasters (P. purpurascens a single verticillaster), normally dense or with lower verticils laxly Stem solid, terete; hairs 7-celled, c. 600 jam long. Leaves ovate, 65 x arranged, rarely lax throughout, with large, ovate, rarely lanceolate 30 mm, base cuneate, apex acute, margin crenate; hairs 4-celled, c. bracts, commonly lobed, deeply cleft and two-lobed, or toothed, 400 urn long. Petiole 15 mm long; hairs 4-celled, c. 270 urn long. equaling or exceeding the calyx. Calyx tubular-inflated to Inflorescence a terminal spike, c. 50 mm long, dense, with more than campanulate, with 5 rib veins, pinnately-branched in region of teeth, two lateral spikes; hairs 3-celled, c. 500 um long; bracts lanceolate incrassate with lines of elongated cells parallel to ribs and coming to ovate, c. 6-7 mm long, some deeply cleft, two-lobed, c. 6 x 2-3 together at sinuses of teeth; teeth triangular, hairy outside, coarsely mm, hairy. Calyx tubular-inflated, 4.0 x 3.7 mm, 5-veined; teeth and hairy or glabrous within except at tips. Corolla tube usually equal to upper part of tube tomentose with long hairs outside, hairy within, limb. more densely above to sparsely below; teeth ciliate, c. 1.0-1.5 mm long, 0.7-0.8 mm wide at base; outer hairs 7-celled, c. 1 600 um long.

1 . (Burm.f.) Kuntze, Rev. gen. pi. 2: 529 Pogostemon benghalensis Corolla up to 6.2 mm long; lower lip c. 1 .8 x 1 .2 mm; upper lip c. 1 (1891). mm across; central lobe 1.1 x 0.5 mm; lobes hairy outside. Filaments 4a, 14a. Figs inserted more or less equally, at a height of c. 2.7 mm in the tube; filaments c. 5.6-6 mm exserted c. 2.5 Origanum benghalense Burm.f., Fl indica: 128, t. 38 f. 3 (1768). long, portion mm; many long hairs at middle and base of some with a few Pogostemon parviflorus Benth. in Wall., PI. asiat. rar. 1: 31 (1830). simple filaments, hairs. c. 1 lobes c. 1 mm. Disc c. P. intermedius Benth. in Wall., Numer. List: 2327 (1830). moniliform Style mm long; stigma 0.8 mm Nutlets 4, c. 900 x 650 um, P.frutescens Graham, Cat. pi Bombay: 149 (1839), pro parte. long. oblong, puncticulate.

P. Dalzell in Hooker's J. Bot. 2: 336 ( 1 850), pro parte. purpuricaulis DISTRIBUTION. India (Bengal) and Bangladesh. P. plectranthoides auct. pro maj., non Desf. SPECIMENS EXAMINED. Hooker s.n. (K), Roxburgh s.n. (BM- Stem solid, angular; hairs 5-celled, c. 460 urn long. Leaves ovate, syntype), Wallich 153 (K-syntype). 130 x 60 mm, base cuneate, apex acuminate, margin double dentate; This is similar to but has smaller hairs 4-celled, c. 330 urn long. Petiole 25 mm long; hairs 4-celled, c. species Pogostemon benghalensis flowers and a tube which is within. 360 urn long. Inflorescence a terminal spike, c. 70 mm long, with calyx hairy 90 G.R. BHATTI AND M. INGROUILLE

Fig. 4 Dissected corollas, calyces, and bracts, showing inner surface. Pogostemon subgenus Pogostemon: (a) P. benghalensis, (b) P. villosus, (c) P. plectranthoides, (d) P. pubescens, (e) P. gardneri, (f) P- cristutus, (g) P. paniculatus. Scale bar 5 mm. SYSTEMATICS OF POGOSTEMON 91

3. Pogostemon plectranthoides Desf. in Mem. Mus. Hist. nat. filament c. 6.5 mm long, exserted portion c. 3.2-4.4 mm; filaments Paris 2: 155, pi. 6 (1815). glabrous towards the base. Style 7.0-9.7 mm long; stigma lobes c. Figs4c, 14c. 0.6-0.8 mm. Disc c. 0.6-0.9 mm long. Nutlets 4, c. 400-500 x 400- 437 um, obovoid, puncticulate to reticulate-foveate. Mentha secunda Roxb., Hort. bengai: 44 (1814). DISTRIBUTION. Southeast Asia (Thailand and Vietnam). Shrubby plant up to 3 m high; stem solid, angular; hairs 4-celled, c. 560 um long. Leaves ovate, 152 x 70 mm, base rounded, apex acute, SPECIMENS EXAMINED. Kerr 2384 (BM), Kerr 3113 (BM), Petelot margin double dentate; hairs 4-celled, c. 500 urn long. Petiole 32 mm 5297 (BM). long; hairs 5-celled, c. 500 urn long. Inflorescence a terminal spike, This species is like Pogostemon plectranthoides but has smaller c. 60 mm long, dense, with more than two lateral spikes; hairs 5- flowers. celled, c. 780 urn long; bracts broadly ovate, c. 6 x 2-4 mm, hairy. Calyx tubular-inflated, 4.8 x 4.4 mm, 5-veined; with short hairs 5. Pogostemon cristatus Hassk. in Hoeven & de Vriese, Tijdschr. outside, only teeth hairy within; teeth ciliate, c. 1.1-1.3 mm long, Natuurl. Gesch. Physiol. 10: 127 (1843). 0.9-1.0 mm wide at base; outer hairs 4-celled, c. 650 um long. Figs 4f, 15c. Corolla up to 8 mm long, purple; lower lip c. 2.2 x 1.1 mm; upper lip c. 2.3 mm across; central lobe 1 .5 mm long, 0.6 mm wide at base. Stem weak, contracted at the base of the internodes, angular, sparsely Filaments purple, inserted at different heights, the lowest at a height hairy; hairs 4-celled, c. 370 u,m long. Leaves ovate, 120 x 80 mm, of 3.5 mm in the tube; longest filament c. 8.0-8.6 mm long, exserted base cuneate, apex acute, margin inciso-crenate; hairs 4-celled, c. portion c. 4. 1 mm; filaments more or less glabrous towards the base. 562 |im long, some covered with a crystalline powder. Petiole c. 60 Style c. 10 mm long, purple; stigma lobes c. 1.2 mm. Nutlets 4, c. mm long; hairs 4-celled, c. 370 um long. Inflorescence a terminal 600 x 450 um, obovoid, puncticulate. spike, c. 100 mm long, lax below and dense above, with more than two lateral spikes; hairs 3-celled, c. 430 (am long; bracts ovate, up to DISTRIBUTION. Throughout the Indian subcontinent from Tamil 6x3 mm, hairy. Calyx tubular, 6x5 mm, 5-veined; hairy outside Nadu to Nepal and Bangladesh; absent from Sri Lanka. A very with long hairs and shorter hairs on veins and margin, teeth and common constituent of the ground flora of Terminalia woodland or upper part of tube hairy within; teeth ciliate, 1.7-1.9 mm long, 0.9- open bush. 1 .0 mm wide at base; outer hairs 4-celled, c. 750 urn long. Corolla up

ETHNOBOTAN Y. Pogostemon plectranthoides is one of the patchouli to 1 1 mm long; lower lip 2.2 x 1.5 mm; upper lip c. 2 mm across, species, called 'Rudilo' in Nepal, 'Thekkali' in India, and used hairy outside; central lobe c. 1.7 x 0.7 mm. Filaments attached at a medicinally. height of 4 mm in the tube, c. 10-1 1 mm long, exserted portion c. 4 mm; filaments glabrous towards the base. Style c. 12 mm long; SPECIMENS EXAMINED. Barber 1353 (K), Bista 2591 (BM), stigma lobes 1 .3 x 1 .6 mm. Disc c. 0.5 mm long. Nutlets 4, c. 900 x Chatterjeo (K), Codrington 380 (BM), Cooper 4991 (K), Dobremez 700 um, oblong, tan, with rounded patches on the surface. 2149 (BM), Drummond 20646 (K), Drummond 26164 (K), Drummond 26165 (K), Drummond 26166 (K), Duthie 10531 (K), DISTRIBUTION. Scattered throughout the East Indies including the Duthie 22399 (K), Falconer (K), Gamble 9007 (K), Gamble 9173 Lesser Sunda Islands and Timor. (K), Gamble 10212 (K), Gamble 13941 (K), Gamble 15843 (K), SPECIMENS EXAMINED. Kostermans & Wirawam 55 1 (K). Gamble 24539 (K), Gardner 230 (BM), Lai s.n. (K), Lambert 1030

1 1 1 van den Brink Jr. who (K), Meebold 4699 (K), Mooney 673 (K), Mooney 2 7 (K), Mooney Keng ( 978: 356) quoted Backer & Bakhuizen 2635 (K), Nanak 20248 (K), Nanak 202647 (K), Polunin, Sykes & had noted that Pogostemon cristatus is closely related tof! heyneanus, Williams 3784 (BM), Rich 508 (K), Ritchie 543 (K), Roxburgh s.n. but may be distinguished by its larger flowers. (BM), Stainton 13 (BM), Stainton 5220 (BM), Stewart 1000 (K), Fl. Brit. India 4: Strachey & Winterbottom 1 (BM), Talbot 1839 (K), Thomson 527 6. Pogostemon gardneri Hook.f., 632 (1885). (K), Verma 6891 (K), Wallich 1530 (K), Wood s.n. (K). Figs4e, 15b.

reddish hairs This species is similar to Pogostemon benghalensis. Differences are Stem erect, bluntly 4-angled, brown, villous; 5-celled, 80 x base listed under that species. c. 1500 (am long. Leaves ovate, 50 mm, cuneate, apex acuminate, margin inciso-crenate or sharply double crenate; hairs 5- celled, c. 1300 (am long. Petiole c. 30 mm long; hairs 6-celled, c. 4. Pogostemon pubescens Benth. inA. DC., Prodr. 12: 152 (1848). 1300 (am long. Inflorescence a terminal spike, c. 80 mm long, dense, Figs4d, 15a. with two lateral spikes, flowers in whorls; hairs on stalk 5-celled, c. Stem solid, terete; hairs 3-5 celled, c. 375-1300 um long. Leaves 1250 um long; bracts ovate, up to 6 x 1.5 mm, hairy. Calyx tubular- ovate, 70-90 x 31-67 mm, base truncate or cuneate, apex acute, inflated, 4 x 4.5 mm, 5-veined; outside with relatively few, stout margin serrate or double crenate; hairs 3-4-celled, c. 850 jam long. hairs, upper half of tube hairy within with long fine hairs, the teeth Inflores- Petiole 20 mm long; hairs 3-5-celled, 475-850 um long. glabrous within; teethe. 0.9 x 0.9 mm; outer hairs 10-celled,c. 1750

than two 1 .5 x 1 cence a terminal spike, c. 45-58 mm long, dense, with more urn long. Corolla up to 5.5 mm long; lower lip mm; upper lip

bracts ovate- 1 all inserted lateral spikes; hairs 3-4-celled, c. 350-480 (am long; c. 1.7 mm across; central lobe c. x 0.5 mm. Filaments x lanceolate, 3-4 x 1-2 mm, hairy. Calyx tubular-inflated, 4.2^.5 at a height of 2 mm in the tube, c. 4.3^.7 mm long; exserted portion with 3.5^.5 mm, 5-veined; hairy outside, teeth and upper part of tube c. 1.2 mm; filaments glabrous at base. Style c. 5.5 mm long; stigma short fine hairs within; teeth ciliate, shortest tooth c. 1.3 x 0.9 mm, lobes 0.5 mm. Disc c. 0.8 mm long. Nutlets 4; c. 1000 x 900 um, urn longest tooth c. 1 .4-1 .6 x 1 .0 mm; outer hairs 3-celled, 560-670 ellipsoid to ellipsoid-oblong, dark brown, puncticulate.

c. 1 .2-1 .4 x 0.5-1 . 1 long. Corolla up to 5.8-6.5 mm long; lower lip DISTRIBUTION. Western India in the vicinity of Travancore. lobe c. 1 0.5- mm; upper lip c. 1 .4-2.3 mm across; central mm long, less at SPECIMENS EXAMINED. Gardner 1 847 Gardner 2 1 22 0.6 mm wide at base, hairy. Filaments inserted more or equally (K-syntype), shortest Venkabo 3229 1913 (K). a height of 2.5 mm in the tube; longest filamentc. 7 mm long, (CAL-syntype), (K), Wight 92 G.R. BHATTI AND M. INGROUILLE

7. Pogostemon paniculatus (Willd.) Benth. in Wall., PI. asiat. rar. SPECIMENS EXAMINED. Dalzell 5747 (K-syntype), Duthie 10537 1:30(1830). (K), Hooker 2337 (K-syntype), Stocks s.n. (BM), Watt 5078 (K), Figs4g, 16a. Wight s.n. (K).

Hyssopus cristatus Lam., Encycl. 3: 187 (1789), non Elsholtzia 9. Pogostemon cablin (Blanco) Benth. in A. DC., Prodr. 12: 156 cristatus (Willd. )Willd. ( 1 800), non Pogostemon cristatus Hassk. (1848). (1843). Figs 5b, 16c. Elsholtzia paniculata Willd., Sp. pi 3: 59 (1800). Mentha cablin Blanco, Fl. Filip.: 473 (1837). Stem solid, angular; hairs 4-celled, c. 750 (am long. Leaves ovate, M. auricularia Blanco, Fl Filip. ed. 2: 473 (1845), non L. (1767). 106 x 57 mm, base cuneate, apex acute, margin double dentate; hairs Pogostemon patchouly Pellet, in Mem. Soc. Sci. phys. Orleans 5: 3-celled, c. 600 u,m long. Petiole c. 1 8 mm long; hairs 5-celled, c, 277, t. 7(1845). 800 u.m long. Inflorescence a terminal spike, c. 40 mm long, secund, P. suavis Ten. in Parl., G. Bot. ital 2: 56 (1847), pro parte. lax below and dense above, with more than two lateral spikes; hairs c. 4-celled, c. 750 jam long; bracts broad, toothed or entire, up to 7 x Stem solid, angular; hairs 4-celled, 680 um long. Leaves ovate, c. 80 3.5 mm. Calyx infundibular, 4.0 x 3.2 mm, 5-veined; hairy outside x 60 mm, base cuneate, apex obtuse, margin dentate; hairs 5-celled, especially on the teeth, teeth and upper tube hairy within; teeth c. 1250 um long. Petiole c. 26 mm long; hairs 5-celled, c. 900 um ciliate, c. 0.9-1.0 mm long, c. 0.4-0.7 mm wide at base; outer hairs long. Inflorescence a terminal spike, c. 45 mm long, lax below and

5-celled, c. 1 100 urn long. Corolla up to 5.9 mm long; lower lip c. 1 dense above, with more than two lateral spikes; hairs 3-celled, c. 150 x 1 mm; upper lip c. 1.5 mm across, hairy; central lobe c. 0.9 x 0.5 um long; bracts ovate to lanceolate, entire or toothed, 4-9 x 0.7-4.0 mm. Filaments inserted at different heights, the lowest at a height of mm. Calyx tubular-inflated, c. 4.5 x 2.8 mm, 5-veined; densely hairy 2.5 mm in the tube; filaments c. 4.5-5.2 mm long, exserted portion outside, teeth and upper part of tube more or less glabrous within, c. 1.8 mm; filaments glabrous towards the base. Style c. 6.3 mm with only a few scattered hairs; teeth ciliate, c. 1.8-1.9 mm long, c. long; stigma lobes c. 0.7-0.8 mm. Disc c. 0.8 mm long. Nutlets 4, c. 0.6 mm wide at base; outer hairs 3-celled, c. 400 um long. Corolla up 500 x 400 um, ellipsoid to ellipsoid-oblong, puncticulate. to 6.2 mm long, mauve; lower lip c. 1.4 x 0.9 mm; upper lip c. 1.1 mm across, slightly hairy; central lobe 1.3 x 0.4 mm. Filaments all DISTRIBUTION. India (Madras, Bombay, Nilgiri, Kannoth, Kunda inserted at a height of 2 mm in the tube, c. 5.8-6.0 mm long, exserted Hills, Travancore, Mysore, Malabar, Carnatic, Salem, Namakkal portion c. 1.8 mm; filaments glabrous towards the base. Style c. 5.8 Dist.) and Thailand. mm long; stigma lobes 1 .4-1 .9 mm. Disc 0.8 mm long. Nutlets 4, c. SPECIMENS EXAMINED. Beddome s.n. (BM), Fernandes 1786 (K), 800 x 600 um, ellipsoid to ellipsoid-oblong, reticulate-foveate. Gamble 9576 (K), Gamble 1 1987 (K), Gamble 15566 (K), Hooker DISTRIBUTION. Widespread in the tropics including Thailand, Fiji, & Thomson s.n. (K), Kerr 17609 (BM), Miguel 249 (K), Perumal Philippines (Luzon), Sumatera, India, Mauritius, and introduced to 21757 (K), Rungachari s.n. (K), Saldanha 15771 (K), Wallich 1561 Florida. This is one of the species which yields patchouli oil and it (K-neotype), Wight 21 19 (K), Wight 2126 (BM, K), Wight 2529 (K). has evidently been introduced into many places. According to

Merrill ( 1 944) it is commonly found in cultivation in the Philippines, 8. Dalzell in Hooker's J. Bot. 2: Pogostemon purpurascens 337 although not on a commercial scale. It flowers in the Philippines but (1850). rarely elsewhere. This fact does not support the suggestion by 16b. Figs 5a, Merrill (1912) that it is not truly native in the Philippines.

Stem solid, terete; hairs 1 3-celled, c. 1500 jam long. Leaves ovate, 100 ETHNOBOTANY. The leaves and stems of Pogostemon cablin are x 68 base double hairs 8- mm, cuneate, apex acute, margin dentate; employed as an insecticide against cockroaches, moths, and other c. 1700 u,m Petiole 25 hairs 1600 celled, long. mm long; 7-celled, um insects and also as a repellent for leeches. In the Philippines the long. Inflorescence a single terminal spike, 80 mm long, dense; hairs 4- crushed leaves are sometimes used for washing. Leaves are also c. 860 bracts ovate to or celled, um long; palmate, obscurely markedly added to baths for their antirheumatic property and an infusion of to 5-6 x 1 .5-5 6.0 x 4.5 5- toothed, up mm, hairy. Calyx tubular, mm, fresh leaves is taken internally to allay painful menstruation. The veined; hairy outside with long hairs, teeth and tube less densely hairy inflorescence of P. cablin is sold in Goa to dress the hair of women. within; teeth ciliate, c. 1 .3-2.2 mm long, c. 0.7-1 . 1 mm wide at base; SPECIMENS EXAMINED. Alston 14403 (K), Clemens s.n. (BM), outer hairs 4-celled, c. 930 um long. Corolla up to 5.5 mm long; lower Degener & Ordonez 14018 (K), Dupont 1769/23 (K-cult.), Kerr lipc. l.lxl mm; upper lip c. 1 mm across; central lobe c. 1 x 0.4 mm, 17069 (K), Merrill 112 (BM, K), Plowman 13285 (K-cult.), Reillo sparsely hairy. Filaments all inserted at a height of 2.4 mm in the tube, 19292 (BM). c. 5.7-7.0 mm long, exserted portion c. 3.9 mm; filaments glabrous towards the base. Style c. 7.2 mm long; stigma lobes c. 2.0-2.5 mm. 10. Pogostemon nepetoides Stapf in Bull misc. Inf. R. hot. Gdns, Disc c. 0.5 mm long. Nutlets 4, c. 500 x 400 jam, ellipsoid to ellipsoid- Kew [1908]: 116(1908). oblong, reticulate with punctate walls. Figs 5c, 17a. DISTRIBUTION. Throughout India (Assam, Himalaya (West), Stem solid, angular; hairs 4-celled, c. 980 um long. Leaves ovate, 91 Concan, Manipur, Bombay). x 64 um, base truncate, apex obtuse, margin inciso-serrate; hairs 4- ETHNOBOTANY. Pogostemon purpurascens contains an astringent celled, c. 1200 um long. Petiole 34 mm long; hairs 4-celled, c. 1 100 resin, an alkaloid, a yellow varnish of a slightly bitter taste and um long. Inflorescence a terminal spike, c. 105 mm long, lax below mouse-like odour of trimethylamine, and a volatile oil with an odour and dense above, with more than two lateral spikes; hairs 3-celled, c. similar to cedar wood. The stem and leaves are a stimulant and 360 um long; bracts ovate to deeply cleft, three-lobed or toothed, 4- styptic. Fresh leaves are used to clean wounds and provide healthy 8 mm long, hairy. Calyx tubular, 4.6 x 3.6 mm, 5-veined; teeth and granulation. In parts of India the roots are reputed to be a remedy for upper part of tube with long hairs outside, within the teeth hairy, the snake bite. tube more or less glabrous; teeth ciliate, 1 .8-2.0 mm long,c. 0.7 mm SYSTEMATICS OF POGOSTEMON 93

Fig. 5 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Pogostemon: (a) P. purpurascens, (b) P. cablin, (c) P.

nepetoides, (d) P. heyneanus, (e) P. nelsonii, (f) P. glaber. Scale bar 5 mm. 94 G.R. BHATTI AND M. INGROUILLE

x 68 base wide at base; outer hairs 2-celled, c. 560 |jm long. Corolla up to 6.2 Stem solid, terete, glabrous. Leaves ovate, 104 mm,

1 c. 1.6 central double lamina Petiole mm long; lower lip c. 1.6 x mm; upper lip mm across; cuneate, apex acute, margin dentate, glabrous. Inflorescence a terminal c. 43 mm lobe 1 .2 x 0.5 mm, with a few hairs. Filaments inserted at slightly 14 mm long, glabrous. spike, lateral all bracts ovate, different heights, the lowest at a height of 2.6 mm in the tube; long, with two spikes, spikes dense, glabrous; 5.0 filaments c. 5.3-6.8 mm long, exserted portion c. 3.2 mm; filaments entire or toothed, up to 8-9 x 2-6 mm, glabrous. Calyx tubular,

1 x with short hairs glabrous towards the base. Style c. 7.2 mm long; stigma lobes .9- 3.8 mm, 5-veined; very outside, glabrous within; teeth 1 .2-1 .4 0.7-0.8 wide at base. Corolla 2. 1 mm. Disc c. 0.9 mm long. Nutlets 4, c. 600 x 400 urn, ellipsoid ciliate, mm long, mm up

to 8.2 lower c. 2.3 x 1 c. 2 mm to ellipsoid-oblong, puncticulate. mm long; lip mm; upper lip across; at central lobe 1 .5 x 0.7 mm, with a few hairs. Filaments inserted DISTRIBUTION. The Philippines (Luzon). different heights, the lowest at a height of 3.5 mm in the tube; SPECIMENS EXAMINED. Micholitz s.n. (K-holotype), Ramos 22432 filaments c. 6.5-7.5 mm long, exserted portion c. 2.8 mm; monili- (BM). form hairs poorly developed, filaments glabrous towards the base. Style c. 10.2 mm long; stigma lobes c. 1.1-1.3 mm. Disc c. 0.6 mm 1 1. Benth. in Wall., PL asiat. rar. 1: 31 Pogostemon heyneanus long. Nutlets 4, c. 600 x 300 urn, oblong, foveolate. (1830). DISTRIBUTION. Southern India (Kerala). FigsSd, 17b. SPECIMENS EXAMINED. Nelson s.n. (BM). Origanum indicum Roth, Nov. pi. sp.: 265 (1821). Pogostemon patchouly sensu Hook.f., Fl. Brit. India 4: 633 (1885), 13. Benth. in PI. asiat. rar. 1: 31 nonPelletier(1845). Pogostemon glaber Wall., (1830). 18a. P. suavis Ten. in Par!., G. Bot. /to/. 2: 56 (1847), pro parte. FigsSf,

Perilla D. Don, 115 (1825). Stem solid, angular; hairs 4-celled, c. 500 urn long. Leaves ovate, 85 polystachya Prodr.fl. nepal: x 45 mm, base cuneate, acuminate, margin double dentate; apex Bush up to 2 m high; stem dark red-purple; hairs 3-celled, c. 250 urn hairs 4-celled, c. 460 u.m long. Petiole 32 mm long; hairs 4-celled, c. long. Leaves ovate, 150 x 97 mm, base cuneate, apex acuminate, 350 urn Inflorescence a terminal c. 70 mm long, lax long. spike, margin double dentate; hairs 3-celled, c. 300 urn long. Petiole 40 mm below, dense above, with more than two lateral spikes; hairs 4- long, glabrous. Inflorescence a terminal spike, c. 55 mm long, dense, celled, 350 urn bracts ovate to lanceolate, entire or indistinctly long; with two lateral spikes; hairs 3-celled, c. 230 urn long; bracts toothed, 3.0-5.0 x 0.8-3.0 mm. tubular, 4.3 x 3.1 mm, 5- Calyx lanceolate, 3.0-7.0 x 0.8-1.3 mm, hairy. Calyx tubular-inflated, c. veined; teeth and of tube within, upper part hairy hairy throughout 3.3 x 3.5 mm, 5-veined; outside the tube sparsely hairy and the teeth outside; teeth ciliate, c. 1.2-1.4 mm long, 0.4-0.6 mm wide at base; glabrous, the teeth hairy within; teeth ciliate, c. 0.9-1 . 1 mm long, c. outer hairs 3-celled, c. 312 urn Corolla to 4.8 mm long. up long; 0.6-0.8 mm wide at base. Corolla up to 4.7 mm long, pale whitish lower 1.5 mm central lobe 0.9 lipc. 1.6x0.9mm;upperlipc. across; 1 pink or light mauve; lower lip c. 1 .2 x 1 mm; upper lip c. .6 mm x 0.3 mm, Filaments inserted more or less equally, at a height hairy. across, hairy; central lobe 0.9 x 0.6 mm. Filaments inserted at of 1.7 mm in the tube, c. 3.7^.3 mm long, exserted portion c. 1.2 different heights, the lowest at a height of 2 mm in the tube; filaments mm; filaments towards the base. Style c. 5.6 mm long; glabrous c. 4.7-5.8 mm long, exserted portion c. 3.1 mm; filaments glabrous lobes c. 0.6-0.7 mm. Disc c. 0.7 mm long. Nutlets 4, c. 500 stigma towards the base. Style 6.5 mm long; stigma lobes 0.9-1 mm. Disc x 400 urn, orbicular, reticulate and punctate. c. 0.7 mm long. Nutlets 4, c. 700 x 600 urn, orbicular, puncticulate. DISTRIBUTION. in southern Asia from Sri Lanka, In- Widespread DISTRIBUTION. Widespread from northwestern India and Nepal to dia to Indonesia (Bombay, Mysore, Tinnually Ghats) (Java, Yunnan and Thailand. In Thailand it is found in mixed-oak forest, in Sumatera). introduced into some areas like the Seychelles, Perhaps shade, above 1000 m, and in Nepal it is a common weed on cut-over although also found beside pathways through the jungle. slopes. ETHNOBOTANY. One of the most sources of oil important patchouli ETHNOBOTANY. The leaves ground in water are used to relieve the which is extracted from the dried tops. It is very rich in volatile pain and itching of mosquito bites. terpenoids.The dried leaves and tops of plants are sold in the markets SPECIMENS EXAMINED. Dawson 262 Dobremez 739 of India and the Seychelles. The leaves of Pogostemon heyneanus (BM), (BM), Flatt 161 Forrest 16161 Gamble 3846 (K), (var. 'Patch-Pat') are used medicinally for stomach and skin diseases (BM), (BM, K), Henry 10418 11487 12832 H.I. 1533 and in Malaysia to treat coughs and asthma. A decoction made from (K), Henry (K), Henry (K), (K- Kerr 1 684 Kerr 2907 Kerr 3922 Kerr the roots is sometimes administered for dropsy. Hartwell (1982) syntypes), (BM), (BM), (BM), 4737 McClure 8454 Nicolson 2941 749 listed P. heyneanus as a possible anti-cancer medicine. In different (BM), (K), (BM), Schilling parts of India the leaves are used as a diuretic and carminative agent (K). and they are generally given with the seeds of Ocimum tenuiflorum 14. Pogostemon hispidusPrain in Bull. misc. Inf. R. hot. Gdns, Kew (vernacular name 'Tulsi') in cases of scanty urine and biliousness. [1908]:254(1908). SPECIMENS EXAMINED. Barber 2218 Beddome s.n. (K), (BM), Figs6a, 18b. Farnandas 1886 (K), Gardner 660 (BM, K), Johnson 130 (K), Stem solid, hairs 6-celled, c. 720 jam long. Leaves ovate, c. Kuntze 5258 (K), Macrae 254 (BM, K), Macrae 1532 (K-type), angular; 1 18 x 87 mm, base cuneate, acute, double dentate; hairs Macrae 1733 (K), Ridley 14300a (BM), Robinson & Kloss 88 (BM), apex margin 4-celled, c. 770 urn Petiole 46 mm hairs 5-celled, 870 jam Simpson 9513 (BM), Wallich 231 (K), Wallich 1830 (K), Wallich long. long; Inflorescence a terminal c. 70 mm dense, with two 1891 (BM), Wallich 2131 (BM), Wight 2120 (K). long. spike, long, lateral spikes; hairs 5-celled, c. 780 urn long; bracts ovate or 12. Pogostemon nelsonii Doan in Humbert, Fl. gen. Indo-Chine 4: lanceolate, indistinctly toothed, 5.0-7.0 x 1.0-1.8 mm, hairy. Calyx 975(1936). tubular-inflated, 4.0 x 3.5 mm, 5-veined; tube more or less glabrous inside the Figs 5e, 17c. within, the teeth very hairy at tip, more sparsely below, SYSTEMATICS OF POGOSTEMON 95

6 Dissected Fig. corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Pogostemon: (a) P. hispidus, (b) P. wattii, (c) P. championii,

(d) P.formosanus, (e) P. elsholtzioides, (f) P. tuberculosus, (g) P. griffithii, (h) P. dielsianus. Scale bar 5 mm. 96 G.R. BHATTI AND M. INGROUILLE

c. Nutlets c. 780 x 650 reticulate- outside with relatively few long hairs throughout; teeth ciliate, long. 4, um, orbicular, black, shiny, 0.8-1.0 mm long, 0.5-0.9 mm wide at base; outer hairs 3-celled, c. foveate. 680 urn Corolla up to 4.5 mm long; lower lip c. 1 .2 x 1 .2 mm; long. DISTRIBUTION. Hong Kong. 1 central lobe c. 0.9 x 0.8 with a few upper lip c. .9 mm across; mm, 12421 hairs. Filaments inserted at different heights, the lowest at a height of SPECIMENS EXAMINED. Champion 339 ( K-holotype), Hu (K). 2 mm in the tube; filaments c. 3.7-4 mm long, exserted portion c. 1 .5 in Hooker's Icon. PI. 25: mm; filaments glabrous towards the base. Style c. 8.5 mm long; 17. Pogostemon formosanus Oliver pi. c. 700 x 600 stigma lobes c. 0.6 mm. Disc c. 0.4 mm long. Nutlets 4, 2440(1896). urn, orbicular, foveate. Figs6d, 19b.

DISTRIBUTION. Thailand to Bangladesh. In forest areas. Stem solid, terete, sparsely hairy; hairs 3-celled with longer apical cell, c. 250 um long. Leaves ovate, 1 1 x 6.5 mm, base cuneate, apex SPECIMENS EXAMINED. Jenkins in Herb. Hooker (K-paratype), acuminate, margin inciso-serrate, abaxial surface of lamina gla- terr 6631 (BM). brous, adaxial surface sparsely hairy; hairs 2-celled, c. 250 um long. Petiole 45 mm long, weak; hairs 2-celled, c. 160 um long. Inflores- 15. Pogostemon wattii C.B. Clarke in J. Linn. Soc. 25: 59 (1889). cence a terminal spike, c. 45 mm long, lax, with three or more lateral Figs6b, 18c. spikes; hairs 2-celled, c. 300 um long; bracts small, lanceolate, c. 3.5 x c. 4 x 3.6 5- Pogostemon battakianus Ridl. in J. Asiat. Soc. Mai. 1: 85 (1923). 0.7 mm, hairy. Calyx campanulate, angular, mm, veined; puberulent outside, mostly on the veins, teeth hairy within Stem solid, terete; hairs 5-celled, c. 800 urn. Leaves ovate, c. 75 x 45 near tip with a row of hairs slightly inside the margin, otherwise mm, base truncate, apex acute, margin double dentate; hairs 5- glabrous; teeth c. 1 .3-1 .5 x c. 0.6-0.8 mm; hairs 2-celled, c. 250 um celled, c. 1000 Petiole c. 20 mm long; hairs 5-celled, c. 0.8 (am long. 1 hairs at long. Corolla c. 4.5 mm long; lower lip c. .2 x 0.9 mm, few mm Inflorescence a terminal spike, c. 1 cm long, dense, with long. 1 central lobe c. 1 .2 x 9 tip; upper lip c. .5 mm across, hairy outside; two or more lateral spikes; hairs 8-celled, c. 1000 (am long. Calyx mm. Filaments all inserted a height of 2.4 mm in the tube, c. 2.5-2.8 tubular-inflated, c. 3 x 3.5 mm, 5-veined; sparsely hairy or glabrous mm long, exserted portion c. 0.7 mm; filaments hairy towards the outside, teeth and upper part of tube hairy within; teeth ciliate, c. base, two upper stamens more so than lower two. Style c. 6 mm long; 0.8-1 mm long, c. 0.5-0.7 mm wide at base; outer hairs 3-celled, c. stigma lobes c. 0.5 mm. Disc c. 0.5 mm long. Nutlets 4, c. 700 x 600 350 urn Corolla up to 4 mm long; lower lip c. 1 x 1 mm; upper long. um, obovoid, dark brown, reticulate-foveate. 0.6 wide at lip c. 1.8 mm across; central lobe c. 0.8 mm long, mm base. Filaments all inserted at a height of 1.7 mm in the tube, c. 3.5- DISTRIBUTION. Taiwan. 4 mm long, exserted portion c. 0.5 mm; filaments all glabrous at the SPECIMENS EXAMINED. Henry 1 178 (CAL, K). base. Style 5.5 mm long; stigma lobes c. 0.5 mm. Disc c. 0.4 mm Nutlets 4; c. 430 x 360 urn, ovoid, reticulate-foveate. long. 18. Pogostemon elsholtzioides Benth. in A. DC., Prodr. 12: 153 DISTRIBUTION. India (western Assam). (1848). Figs 6e, 19c. SPECIMENS EXAMINED. Clarke 41719 (K-holotype). Bushy shrub up to 2 m high; stem solid, angular; hairs 4-celled, c. The name Pogostemon wattii C.B. Clarke has been a source of 220 um long. Leaves lanceolate, 120 x 28 mm, base cuneate, apex confusion. Prain (1891) included P. wattii in 'Some additional acuminate, margin serrate; hairs 4-celled, 225 um long. Petiole 0.6 species of Labiatae', remarking that it was very distinct. Later he mm long; hairs 4-celled, c. 225 um long. Inflorescence a terminal described the same he had Eurysolen gracilis, citing specimens hairs spike, c. 50 mm long, dense, with more than two lateral spikes; previously listed under Pogostemon wattii. It is doubtful if he saw 3-celled, c. 170 um long; bracts ovate or lanceolate, 1.5-5.0 x 0.6- the holotype of P. wattii which is very different. Use of Prain's 1.4 mm. Calyx tubular, 4.2 x 3.0 mm, 5-veined; teeth hairy within, account has resulted in the misidentification of many specimens of hairy outside with many white hairs and with small tufts of hair at Eurysolen gracilis as Pogostemon wattii. sinuses of teeth; teeth not ciliate, c. 1.0-1.1 mm long, c. 0.5-0.7 mm wide at base; outer hairs 3-celled, c. 160 um long. Corolla up to 4.5 16. Pogostemon championii Prain in Bull. misc. Inf. R. hot. Gdns, mm long, purple; lower lip c. 1 x 0.7 mm; upper lip c. 1.7 mm across, Kew [1908]: 254 (1908). glabrous; central lobe 0.7 x 0.9 mm. Filaments pink, inserted at Figs 6c, 19a. different heights, the lowest at a height of 2 mm in the tube; filaments towards Stem erect, solid, 4-angled, glabrous below, hairy upper c. 4.5-5 mm long, exserted portion c. 2.5 mm; filaments with a few

to 1 x 60 parts; hairs 5-celled, c. 500 jam long. Leaves ovate, up 10 hairs, more or less equally hairy towards the base. Style 5.8 mm double hairs 4- mm, base cuneate, apex acuminate, margin crenate; long; stigma lobes c. 0.7 mm long. Disc c. 0.5 mm long. Nutlets 4, celled, c. 550 urn long. Petiole up to 40 mm long; hairs 5-celled, c. c. 1700 x 600 um, lanceolate, ruminate. 500 urn long. Inflorescence a terminal terminal spike, c. 40 mm DISTRIBUTION. Widespread in the Himalayas: Bhutan, India long, first whorl of flowers separated from the rest, lateral spikes (Khasia, Naga Hills). Common in secondary growth, waste places, with single whorl of flowers; hairs on stalk 4-celled, c. 400 urn along roadsides, at the edge of forests, on cultivated land, and around long. Calyx tubular-inflated, c. 4.2 x 3.2 mm, 5-veined; hairy villages. outside especially on veins, glabrous within; teeth ciliate, 1.1-1.3 mm long; outer hairs 4-celled, c. 550 urn long. Corolla up to 5.5 SPECIMENS EXAMINED. Bar 24 (K), Bar 6649 (K), Clarke 4 1 505

mm long; lower lip 1.5 x 1 mm; upper lip 1.2 mm across, hairy; (K), Clarke 42819 (BM), Griffith 202 (BM, K-syntypes), Griffith 6449 central lobe c. 1 x 0.5 mm. Filaments inserted at different heights, 3962 (K-syntype), Kingdon-Ward 6346 (K), Kingdon-Ward the lowest two at a height of 2.5 mm in the tube; filaments c. 5-6 (K), Kingdon-Ward 7841 (K), Kingdon-Ward 11293 (BM, K), mm long, exserted portion c. 2.5 mm; filaments glabrous at the Kingdon-Ward 14234 (BM), Kingdon-Ward 18372 (BM), Kingdon- base. Style c. 7.5 mm long; stigma lobes c. 0.7 mm. Disc c. 0.8 mm Ward 18894 (BM), Kingdon-Ward 20274 (BM), Ludlow, Sherriff & SYSTEMATICS OF POGOSTEMON 97

Elliot 12255 (BM), Meebold 17.021 (K), Simon s.n. Watt 6656 (K), Spreading shrub up to 3 m; stem solid, angular; hairs 3-celled, c. 100 (K), um long. Leaves linear-lanceolate, 120 x 32 mm, base cuneate, apex acuminate, margin dentate, lamina more or less glabrous above, with 19. Pogostemon tuberculosus Benth. in Wall., PL asiat. rar. 1: 31 reddish 3-celled hairs below, c. 150 um long. Petiole c. 1 mm long; (1830). hairs 3-celled, c. 125 um long. Inflorescence a terminal spike, c. 45 6f, 20a. Figs mm long, dense, with more than two lateral spikes; hairs 3-celled, c. 1 80 um long; bracts lanceolate, c. 4.5 x 0.8 mm, hairy. tubular, Shrub 2-3 m high; stem solid, terete; hairs stellate, central hair 2- Calyx c. 4.4 x 3.0 mm, 5-veined; teeth hairy within with long fine hairs, a celled, c. 1 10 |am long, radiating hairs 1 -celled. Leaves ovate, 1 10 x row of hairs slightly inside the margin, outside with and a 65 mm, base cuneate, apex acute, margin crenate; hairs similar to glands dense felt of short hairs; teeth c. 1 .2 x 0.7 mm; outer hairs 3-celled, those of stem. Petiole 30 mm long; hairs similar to those of stem. c. 1 25 um Corolla to long. up 7.5 mm long, rose-red; lower lip 1 .4 x Inflorescence with more than two laterals verticillasters; verticillasters 1 mm; upper lip c. 1.8 mm across; central lobe c. 1.2 x 1 mm. c. 200-300 mm long; verticils many flowered, very dense, widely Filaments inserted at different the at a in heights, lowest height of 3 mm spaced inflorescence; bracts small, ovate, 2.5 x 1.0 mm, hairy. in the tube; filaments c. 6.2-7 mm long, exserted portion c. 3.5 mm; Calyx tubular-inflated, 3.0 x 3.0 mm, 5-veined; with many stellate two filaments villous towards the base. c. 6.4 hairs Style mm long; stigma outside, teeth hairy within; teeth ciliate, c. 0.9-1 .0 mm long, c. lobes c. 1 .2 mm. Disc c. 0.7 mm long. Nutlets 4, c. 1500 x 800 um, 0.4-0.6 mm wide at base; outer hairs with central hair 3-celled, c. lanceolate, ruminate. urn 1 370 long, lateral hairs -celled. Corolla up to 7.2 mm long, pale

lower c. 1 .5 x 1 c. 2 mauve; lip mm; upper lip mm across, glabrous; DISTRIBUTION. Western China (Yunnan, Salween Irrawaddy di- central lobe c. 1 .3 x 0.8 mm. Filaments all inserted at a height of 2.4 vide, and valley of the Salween). Amongst rocky scrub on dry rocky mm in the tube, deep purplish, c. 7.4-8.9 mm long, exserted portion hillsides. c. 4. 1 mm; filaments glabrous at the base, anthers fawn. Style deep SPECIMENS EXAMINED. Forrest 875 (E, K-isotypes), Henry 9082 purplish, 11.5mmlong;stigmalobesc. 12mm. Disc c. 0.7 mm long. (K), Henry 1 1 174 (K), Henry 12563 (K). Nutlets 4, c. 800 x 600 um, oblong, granulose.

DISTRIBUTION. Eastern Himalayas (Bhutan, Nepal, Kaluimphoong, II. Subgenus ALLOPOGOSTEMON Bhatti & Ingr., nov. Assam, Darjiling). Among thick undergrowth in broad-leaved forest. subgen. section Racemosa SPECIMENS EXAMINED. Clarke 26368B (K), Gamble 3843A (K), Pogostemon Benth., pro parte. Gamble 3847A (K), Gamble 7515 (K), Haines 1041 (K), Hooker Inflorescentia plerumque simplex. Bracteae lanceolatae, lineares s.n. (K), Ludlow, Sherriff& Taylor 61'59 (BM, E), Treutler 1 194 (K), vel filiformis. Calyx tubulosus, actinomorphus, 5 nervi medii plus Wallich s.n. (K-type). usque ad 5 nervos secundarios marginem attingentes, introrsum annulo hirsute. Filamentis a basi villosis vel glabratis, a medium 20. Pogostemon griffithii Prain in Bull. misc. R. hot. Gdns, Kew Inf. pilibus moniliformibus vel glabratis. [1908]: 181 (1908). Inflorescence an unbranched bracts lanceo- Figs 6g, 20b. normally verticillaster, late, linear or filiform, equalling or much smaller than calyx; calyx Pogostemon griffithii var. C.Y. Wu & Y.C. Fl. latifolius Huang, with 5 main rib veins plus isolated branched secondaries reaching Yunnanical: 744(1977). sinuses of teeth, calyx interior glabrous or with a continuous or broken annulus of hairs around of tube and teeth; filament bases Stem solid, terete; hairs 3-celled, 250 um long. Leaves 2, lanceolate, top villous or with or without moniliform hairs. 95 x 25 mm, base cuneate, apex acute, margin dentate; hairs 3- densely glabrous, celled, c. 250 um long. Petiole c. 10 mm long; hairs 3-celled, c. 250 Ha. Section RACEMOSUS (Benth.) Bhatti & stat. um long. Inflorescence a terminal spike, c. 50 mm long, dense, with Ingr., nov. two lateral spikes; hairs 3-celled, c. 250 um long; bracts ovate or 2.0-3.0 x 0.5-1.0 3.3 x lanceolate, mm, hairy. Calyx campanulate, Pogostemon section Racemosa Benth., Pogostemon section Barbata 3.5 mm, 5-veined; of tube and teeth within, upper part hairy hairy Briq., pro parte. throughout outside; teeth ciliate, c. 1.1-1.4 mm long, 0.5-0.7 mm Inflorescence, bracts and bracteoles linear to linear-lanceolate, wide at base, incurved in fruit; outer hairs 3-celled, c. 375 um long. large or small, verticillaster unbranched, or if branched then calyx with 5 Corolla up to 5 mm long; lower lipc. 1 .5 x 1 mm; upper lipc. 1 .2 mm main rib veins plus isolated branched secondaries from sinuses of across; central lobec. 1 .3 x 0.7 mm. Filaments all inserted at a height teeth, with an annulus of hairs around top of tube and teeth, or of 2 mm in the tube, c. 4.7-5.2 mm long, exserted portion c. 2.2 mm; filament bases densely villous. filaments more or less glabrous towards the base. Style c. 5.5 mm long; stigma lobes c. 1.2 mm. Disc c. 0.4 mm long. Nutlets 4, c. 800 Ha i. Subsection RACEMOSUS x 600 um, oblong, ruminate. With moniliform hairs on the filaments. DISTRIBUTION. Burma.

22. Fl. Madras: 1 134 SPECIMENS EXAMINED. Griffith 3962 (K-holotype). Pogostemon nilagiricus Gamble, (1924). Figs 7a, 2 la. The type specimen of Pogostemon elsholtzioides is also Griffith Stem hairs whitish, 12-celled, c. 2500 um 3962, although the collection is of a different plant from a different solid, bluntly terete; long. Leaves ovate, c. 45 x 33 base rounded, acute, locality. mm, apex bluntly margin crenate, with shiny hairs on both sides of lamina; hairs 10- celled, c. 2500 um Petiole c. 10 mm hairs similar to those 21 . Pogostemon dielsianus Dunn in Notes R. hot. Gdn Edinb. 8: 159 long. long; leaves. Inflorescence a terminal to 80 mm (1913). of single spike up long, and hairs on stalk, hairs 10- Figs 6h, 20c. dense; glandular eglandular eglandular 98 G.R. BHATTI AND M. INGROUILLE

Fig. 7 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Allopogostemon section Racemosus subsection Racemosus: (a) P. nilagaricus, (b) P. vestitus, (c) P. mollis, (d) P. rotundatus, (e) P. rupestris, (f) P. rogersii, (g) P. petiolaris. Scale bar 5 mm. SYSTEMATICS OF POGOSTEMON 99 celled, c. 2500 jam long; bracts lanceolate or linear, 4-6 x 0.3-1.3 SPECIMENS EXAMINED. Heyne 1534 (BM, K-syntypes), Heyne mm, hairy. Calyx tubular, c. 4.3 x 3.5 mm, 10-veined; the tube 1561 (BM-syntype), Wight 2126 (K), Wight 2527 (E, K-syntypes). glabrous within, the teeth sparsely hairy, outside hairy; teeth ciliate, c. 1 .5-1 .7 mm long,c. 0.6-0.8 mm wide at base; outer hairs 6-celled, 25. Pogostemon petiolaris Benth. in A. DC., Prodr. 12: 154 (1848). c. 870 urn long. Corolla up to 6 mm long; lower lip c. 2.3 x 1.1 mm; Figs 7g, 23a. upper lip c. 1 .5 mm across; central lobe c. 1 .6 x 0.8 mm; upper and Stem solid, angular; hairs 7-celled, c. 1 100 urn long. Leaves ovate, lower lip hairy outside. Filaments inserted at different heights, the c. 55 x 41 mm, base truncate, apex acute, margin double dentate; lowest at a height of 3 mm in the tube; filaments c. 5.0-5.5 mm long, hairs 6-celled, c. 900 (am long. Petiole 32 mm long; hairs 7-celled, c. exserted portion c. 2.5 mm; filaments thickened towards the base, 1000 urn long. Inflorescence a single terminal spike, c. 75 mm long; the two filaments under upper lip more hairy at the base. Style c. 8 hairs 6-celled, c. 1200 urn long; bracts filiform, 2-3 x 0.1-0.3 mm, mm long; stigma lobes c. 0.5 mm. Disc c. 0.4 mm long Nutlets 4, c. hairy. Calyx tubular, 5.0 x 5.2 mm, 5- veined; with a few long hairs 500 urn, orbicular, dark brown to black, reticulate-foveate with outside, glabrous within, except at the tip of the teeth; teeth ciliate, secondary reticulation. c. 1 .2-1 .5 mm long,c. 0.9-1 .2 mm wide at base; outer hairs 6-celled, DISTRIBUTION. Southern India, Nilagiri (Kunda Hills near c. 1000 um long. Corolla up to 6.5 mm long; lower lip c. 2.5 x 1.2

Ootacamund). mm; upper lip c. 1.6 mm across; central lobe c. 1.7 x 0.7 mm. Filaments inserted at different heights, the lowest at a height of 2 mm SPECIMENS EXAMINED. Bourne s.n. (K-isotypes), Hooker s.n. (K), in the tube; filaments c. 6.2-7.5 mm long, exserted portion c. 3 mm; Rao 46997 (CAL). filaments strigose towards the base, the two upper more densely. Style c. 8.2 mm long; stigma lobes c. 0.7-0.8 mm. Disc c. 1 mm 23. Pogostemon mollis Benth., Labial, gen. spec.: 155 (1833). long. Nutlets 4, c. 600 x 500 um, orbicular, reticulate-foveate with Figs 7c, 21c. secondary reticulation. Stem solid, terete; hairs 6-celled, c. 1750 urn long. Leaves ovate, DISTRIBUTION. Southern India (Western Ghats). soft, c. 32 x 24 mm, base rounded, apex obtuse, margin double dentate; hairs 4-celled, c. 1500 urn long. Petiole c. 8 mm long; hairs SPECIMENS EXAMINED. Hohenacker 1224 (BM, K). 7-celled, c. 2000 urn long. Inflorescence a single terminal spike, c. Labial, 156 55 mm long, lax below and dense above; hairs 6-celled, c. 1000 urn 26. Pogostemon rupestris Benth., gen. spec.: (1833), Dalzell long; bracts linear or filiform, 1.5-7.5 x 0.2-0.9 mm. Calyx in- non Dysophylla rupestris (1851). fundibular, c. 3.8 x 4.0 mm, 10-veined; very hairy outside, glabrous Figs 7e, 22b. within; teeth ciliate, c. 1.3-1.5 mm long,c. 0.7-0.8 mm wide at base; Annual herb with straggling branches; stem angular, swollen and outer hairs 5-celled, c. 410 urn long. Corolla up to 5.5 mm long; rooting at the nodes, glabrous except for a ring of hairs at the nodes. 1 central lower c. 2. 1 x 1.1 mm; upper lip c. .5 mm across, hairy; lip Leaves ovate to ovate-lanceolate, up to 25 x 15 mm, base cuneate to lobe 1.3 x 0.6 mm. Filaments inserted at more or less the same truncate, apex acute, margin dentate towards the apex, lamina height, the lowest at a height of 1 .8 mm in the tube; filaments c. 4.5- sparsely hairy and glandular-punctate; hairs 4-celled, c. 400 um exserted c. 1.3 filaments 5.5 mm long, portion mm; equally strigose lamina. long. Petiole up to 15 mm long; hairs similar to those of at the base. Style c. 6.2 mm long; stigma 1.0-1.2 mm. Disc c. 0.4 mm Inflorescence a single terminal spike, c. 40 mm long, lax; hairs on long. Nutlets 4, c. 700 x 550 jam, orbicular, reticulate-foveate. the stalk 4-celled, c. 400 um long; bracts lanceolate to spathulate, c. x 0.9 tubular to c. 3.5 x 3.8 mm, 10- DISTRIBUTION. India (Bombay, Concan, Western Ghats, 4.0 mm, hairy. Calyx urceolate, veined; within, outside with sparse hairs above; teeth Ootacamund, Nilgiri, Kunda Hills, Pulney Hills). glabrous c. 0.9-1 . 1 x 0.7 unequal, triangular, apex blunt, margins ciliate, mm; Barber 37'62 Bourne SPECIMENS EXAMINED. Anglade 893 (K), (K), outer hairs c. 4-celled, 400 aem long. Corolla c. 4.6 mm long, rose or s.n. 5355 Clarke 10673 Dalzell s.n. (K), Hooker (K), 1 c. 2.8 central (K), (BM), pinkish; lower lip c. 2 x .3 mm; upper lip mm across, Vine 216 2124 2525 at a of c. 2 in (BM), Wight (K-isotype), Wight (K). lobe c. 1 . 1 x 0.9 mm. Filaments all inserted height mm the tube, c. 4.0-5.2 mm long, exserted portion c. 2.6 mm; two 1 rar. 1: 31 staminal hairs 24. Pogostemon vestitus Benth. in Wall.,/ /, asiat. (1830). filaments under central lobe slightly hairy at the base, c. 0.6 Figs7b, 21b. towards the middle blue. Style c. 6.2 mm long; stigma lobes 2 or c. 570 x 500 mm long. Disc c. 0.3 mm long. Mature nutlets 4, Stem solid, terete, very hairy especially at the base; hairs 7-celled, c. um, orbicular, light brown, shiny, reticulate-foveate. 1500 urn long. Leaves ovate, soft felted, 40 x 30 mm, base rounded, Sri Lanka Distr., Nuwara- apex acute, margin dentate; hairs c. 5-celled, c. 1100 urn long. DISTRIBUTION. (Adam's Peak, Kandy in shade on the floor of montane Petiole c. 25 mm long; hairs 5-celled, c. 1200 urn long. Inflorescence Eliya). Growing damp secondary c. 1500 a single terminal spike, c. 30 mm long, dense; hairs 5-celled, forest. urn tubular, 5.8 x 4.8 mm, 10-veined; densely hairy long. Calyx SPECIMENS EXAMINED. Cramer 3800 (K), Cramer 3807 (E), c. 0.7- outside, teeth hairy within; teeth ciliate, c. 0.6-0.8 mm long, Cramer 4277 (K), Cramer 4390 (E, K), Cramer & Jayasuriya 3725 0.9 mm wide at base; outer hairs 3-celled, c. 3200 urn long. Corolla (E), Macrae 396 (K-holotype), Thwaites 343 (BM). c. 2.5 across; c. 8.6 mm long; lower lip c. 3 x 2 mm; upper lip mm at central lobe c. 1.4 x 1.1 mm wide at base. Filaments inserted misc. R. hot. 4.5 in the 27. rogersii N.E. Br. in Bull. Inf. Gdns, different heights in the tube, the lowest at a height of mm Pogostemon c. two 1909]: 379 (1909). tube; filaments c. 7.0-8.5 mm long, exserted portion 1.5 mm, Kew[ 1.2 7f, 22c. filaments tomentose at the base. Style c. 12 mm long; stigmas c. Figs mm. Disc c. 0.4 mm Nutlets 4, c. 600 x 500 urn, oblong, hairs 5- long. Stem procumbent, rooting at the nodes, solid, angular; reticulate-foveate. celled, c. 430 um long. Leaves elliptic to elliptic-oblong, soft-flaccid, 62 x 24 base cuneate, apex acute, margin serrate; hairs 4-celled, DISTRIBUTION. Southern India (western Tamil Nadu). mm, 100 G.R. BHATTI AND M. INGROUILLE c. 450 jam long. Petiole c. 20 mm long; hairs 4-celled, 480 jam long. long; lower lip c. 2.5 x 1.9 mm; upper lip c. 2.3 mm across, hairy; Inflorescence a terminal spike, c. 30 mm long, dense, with more than central lobe c. 1.5 mm long, c. 1 mm wide at base. Filaments inserted two lateral spikes; hairs 5-celled, c. 720 um long. Calyx infundibu- at different heights, the lowest at a height of 4.5 mm in the tube; lar, 2.1 x 2.2 mm, 10-veined; hairy outside, glabrous within; teeth filaments c. 8.4-9.0 mm long, exserted portion c. 4.5 mm; two ciliate, c. 0.7-0.9 mm long, c. 0.4-0.5 mm wide at base; outer hairs filaments villous towards the base. Sty lee. 12 mm long; stigma lobes up to 6-celled, c. 950 urn long. Corolla 6.4 mm long, whitish violet; 1.1-1.2 mm. Disc c. 0.5 mm long. Nutlets 4, c. 400 x 200 um, lower lipc. 1.9x 1.4 mm; upper lip c. 1.7 mm across; central lobe 1.4 oblong, reticulate. x 0.5 mm. Filaments inserted at different heights, the lowest at a DISTRIBUTION. East Indies, Philippines (Panay, Luzon), and height of 3.4 mm in the tube; longest filament c. 5.5-5.7 mm long, Mariana Islands. exserted portion c. 2.7 mm; filaments equally villous at base. Style c. 7.7 mm long; stigmas c. 1.1 x 1.4 mm, violet-purple. Disc c. 0.5 SPECIMENS EXAMINED. Ramos 33320 (BM, K), Ramos & Edano mm long. Nutlets 4, c. 400 x 300 urn, orbicular, reticulate-foveate. 45012 (BM), Santos 31792 (BM), Vidal 1659 (K), Vidal 3421 (K- syntype), Whitehead s.n. (BM-syntype), DISTRIBUTION. Southern Africa (Zimbabwe, Zambia, Angola, South Africa, Mozambique). In marshy ground. 30. Pogostemon velatus Benth. in A. DC., Prodr. 12: 155 (1848). ETHNOBOTANY. Produces edible 'tubers'. Figs 8b, 23c.

SPECIMENS EXAMINED. Fanshawe 3435 (K), Fanshawe 9336 (K), Stem solid, angular; hairs dendromorphic and fruticose, 6-celled, c. Gossweiler 14137 (K), Henriques & Brites 1132 (K), Hundt 149 1250 um long. Leaves ovate, 33 x 19 mm, base rounded, apex (BM), Johnston 983 (K), Multimushii 1023 (K), Rogers 8314 (K- obtuse, margin dentate, hairs dendromorphic and fruticose, 4-celled, holotype), Silva 3801 (K), Stohm 123 (K). c. 1100 um long. Petiole c. 5 mm long; hairs dendromorphic and fruticose, 9-celled, c. 1750 um long. Inflorescence a single terminal According to Brown (1909) this was the first species of Pogostemon spike, c. 165 mm long, dense; hairs dendromorphic; bracts linear or to be reported from Africa and it is quite distinct from Asiatic filiform, 2.5-7.0 x 0.3-0.5 mm, hairy. Calyx tubular, 5.4 x 4.1 mm, species. 10-veined; densely hairy outside, margin of teeth hairy within, otherwise glabrous; teeth ciliate, c. 1 .0-1 .2 mm long, c. 0.8-1 .0 mm 28. Pogostemon rotundatus Benth. in Wall., PL asiat. rar. 1: 31 wide at base; outer hairs dendromorphic, 4-celled, c. 465 um long. (1830). Corolla up to 8 mm long; lower lip c. 2.5 x 1 .4 mm; upper lip c. 2.5 Figs 7d, 22a. mm across, hairy outside; central lobe c. 1.2 x 0.9 mm. Filaments Stem solid, terete; hairs 7-celled, c. 2300 urn long. Leaves orbicular, inserted at different heights, the lowest filaments at a height of 3.8 c. 50 x 25 mm, base truncate to cordate, apex obtuse, margin double mm in the tube; filaments c. 8.0-8.8 mm long, exserted portion c. 4.6 crenate; hairs shiny, whitish, 7-celled, c. 1300 urn long. Petiole up to mm; two filaments strigose towards the base. Style c. 10 mm long; 20 mm long; hairs 7-celled, c. 2000 urn long. Inflorescence a single stigma lobes 1.1-1.2 mm. Disc c. 0.5 mm long. Nutlets 4, c. 500 x terminal spike up to 1 10 mm long, flowers in whorls, arranged side- 300 um, oblong, spinulose. by-side; hairs 7-celled, c. 2000 urn long; bracts lanceolate to filiform, DISTRIBUTION. Philippines (Luzon). 3.5-5.5 x 0.2-1.2 mm. Calyx tubular, c. 6 x 5.5 mm long, 10-veined; very hairy outside, especially the teeth, more or less glabrous within; SPECIMENS EXAMINED. Cuming 1097 (BM, K-isotypes), Elmer teeth ciliate, c. 1.6-1.9 mm long; outer hairs up to 4-celled, c. 900 s.n. (BM), Loher 4209 (K), McGregor 1 1339 (BM), Mendoza 40923 urn long. Corolla c. 9 mm long; lower lip 3 x 1 .8 mm; upper lip c. 3 (BM), Merrill s.n. (BM). mm across, villous outside; central lobe c. 2.4 x 1.5 mm. Filaments inserted at different heights, the lowest at a height of 3.7 mm in the 31. Pogostemon williamsii Elmer in Leafl. Philipp. Bot. 9: 3197 tube; filaments c. 8-9 mm long, exserted c. 3.7 mm; fila- portion (1934). ments widening towards the base, the two upper densely strigose Figs 8c, 24a. towards the base. Style c. 13 mm long; stigma lobes c. 0.5-1 mm. Stem hairs and c. Disc c. 0.5 mm long. Nutlets 4, c. 400 x 300 um, oblong, black, solid, terete; dendromorphic fruticose, 5-celled, covered by more or less circular scales. 600 um long. Leaves lanceolate, c. 105 x 35 mm, base cuneate, apex acuminate, margin crenate; hairs fruticose, 5-celled, c. 800 um long. DISTRIBUTION. Southern India (Madras). Petiole 26 mm long; hairs fruticose, 5-celled, c. 800 um long. Inflorescence a terminal c. 95 hairs SPECIMENS EXAMINED. Heyne s.n. (K-syntype), Wallich 1535 (K- single spike, mm long, dense; c. 8 1 5 bracts lanceolate to 4.0- syntype). fruticose, 5-celled, um long; filiform, 6.0 x 0.2-1.5 mm, hairy. Calyx tubular, 5.0 x 4.3 mm, 10-veined; 29. Pogostemon philippinensis S. Moore in J. Bot., Land. 43: 146 sparsely hairy outside, margin of teeth hairy within; teeth ciliate, c. (1905). 0.6-0.8 mm long, c. 0.7-0.9 mm wide; outer hairs fruticose, 4- 23b. Figs 8a, celled, c. 650 um long. Corolla up to 8 mm long; lower lip c. 2.5 x 1.5 mm; upper lipc. 2.7 mm across, hairy outside; central lobe 1.5 x Stem solid, angular; hairs 4-celled, c. 280 um long. Leaves ovate, 68 0.8 mm wide at base. Filaments inserted at different heights, the x 34 mm, base rounded, apex acute, margin dentate; hairs 5-celled, lowest at a height of 3.5 mm in the tube; filaments c. 10-11 mm long, c. 500 um long. Petiole 20 mm long; hairs 4-celled, c. 440 jam long. exserted portion c. 6.5 mm; two filaments strigose towards the base. Inflorescence a single terminal spike, c. 125 mm long, lax below, Style c. 12 mm long; stigma lobes c. 1.1-1.2 mm. Disc c. 0.6 mm dense above; hairs 5-celled, 600 jam long; bracts lanceolate to long. Nutlets 4, c. 900 x 600 um, ellipsoid to ellipsoid-oblong, filiform, 1.3-4.5 x 0.2-1.5 mm, hairy. Calyx tubular, 5.6 x 5.0 mm, spinulose. 10-veined; hairy outside especially on veins, only teeth hairy within, with a few hairs; teeth ciliate, c. 0.9-1 .0 mm long, 0.9-1 .0 mm wide DISTRIBUTION. Phillippines (Luzon, Mt. Pinatubo, Mt. at base; outer hairs 3-celled, c. 370 um long. Corolla up to 9 mm Semenublam, Baguio, Benguet). SYSTEMATICS OF POGOSTEMON 101

Fig. 8 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Allopogostemon section Racemosus subsection Racemosus:

(a) P. philippinensis, (b) P. velatus, (c) P. williamsii, (d) P. membranaceus, (e) P. elatispicatus. Scale bar 5 mm. 102 G.R. BHATTI AND M. INGROUILLE

base. 13.3 SPECIMENS EXAMINED. Elmer 22225 (BM, K-isotypes), Iwatsuki, c. 4.6 mm; two filaments setose towards the Style mm 0.7 Nutlets c. 330 x Murata & Gutierrez 318 (K), Merrill 1 1673 (BM, K). long; stigma lobes c. 0.6 mm. Disc mm long. 4, 200 urn, semi-circular or 'D' -shaped, spinulose. Keng (1978) and Press (1982) treated Pogostemon williamsii as its and DISTRIBUTION. conspecific with P. velatus but it differs in trichomes, nutlets, Philippines. leaf shape. This taxon has affinities with Pogostemon philippinensis, P. membranaceus, P. williamsii, and P. velatus but it differs by having 32. Pogostemon membranaceus Merr. in Philipp. J. Sci. (Bot.) 7: dendromorphic hairs only on the abaxial surface, setose hairs at the 347(1912). basal part of the filaments, and semi-circular or 'D' -shaped nutlets. Figs 8d, 24b. The verticillaster is very long with a very regular arrangement of

Stem solid, terete; hairs 4-celled, c. 370 um long. Leaves ovate, c. 70 verticils. x 34 mm, base cuneate, apex acute, margin crenate; hairs 3-celled, c. c. 400 34. DC., Prodr. 12: 154(1848). 250 |um long. Petiole c. 14 mm long; hairs 4-celled, um long. Pogostemon paludosusBenth.inA. lax 25a. Inflorescence a single terminal spike c. 140 mm long, below, Figs 9a, dense above; hairs 5-celled, c. 600 jam long; bracts early deciduous. Stem solid, angular; hairs 5-celled, c. 600 urn long. Leaves ovate, c. tubular, 7.0 x 4.0 mm, 10-veined; hairy outside, glabrous Calyx 70 x 35 mm, base somewhat truncate, apex obtuse, margin double c. 1 .2-1 .4 within except at the tip of the teeth; teeth ciliate, mm long, crenate; hairs 5-celled, c. 700 um long. Petiole c. 45 mm long; hairs c. 0.7-0.9 mm wide at base; outer hairs 3-celled, c. 376 (jm long. 5-celled, c. 700 um long. Inflorescence a single terminal spike, c. 90 . 1 2 Corolla up to 8.5 mm long; lower lipc. 2 x 1 mm; upper lipc. mm mm long, lax; hairs 5-celled, c. 1000 um long. Calyx tubular, 3.5 x across; central lobe 1.3 x 0.6 mm. Filaments inserted at different 2.8 mm, 10 veins obscure, 5 veins distinct; annulus of hairs in the in the filaments c. 7.5- heights, the lowest at a height of 4.3 mm tube; calyx throat, outside with long and short hairs; teeth ciliatec. 0.5-0.8 8.8 mm long, exserted portionc. 4.6 mm; filaments equally tomentose mm long, c. 0.4-0.6 mm wide at base; outer long hairs 5-celled, 750 towards the base. Style c. 12.5 mm long; stigma lobes c. 0.5 mm. um long. Corolla up to 5. Omm long; lower lipc. 1.9x 1.5 mm; upper c. 1 Nutlets c. 800 x 300 'D' Disc mm long. 4, (am, -shaped, x inserted lip c. 2.0 mm across; central lobe 0.9 0.4 mm. Filaments reticulate. at different heights, the lowest at a height of 2 mm in the tube;

c. 4.0-4.5 exserted c. 1.5 two DISTRIBUTION. Philippines (Luzon, Laguna, San Antonio). filaments mm long, portion mm; filaments tomentose towards the base. Style c. 6 mm; stigma lobes SPECIMENS EXAMINED. Ramos 16419 (BM-holotype), Ramos 0.8 mm long. Ovary glandular. Discc. 0.5 mm long. Nutlets 4, c. 650 14419 (K), Ramos 16596 (K). x 600 um, orbicular, with cup-like glands.

DISTRIBUTION. India (Nilgiri, Kunda Hills, Tamil Nadu). 33. Pogostemon elatispicatus Bhatti & Ingr., sp. nov. Figs 8e, 24c. SPECIMEN EXAMINED. Klackenberg & Lundin 116 (K).

Herbae hirsutae. Caules subquadrangulati. Folia opposita, lamina 25-75 x 20-35 mm, ovata, crenata, petioli 10-25 mm, abaxialis Ha ii. Subsection GLABRIUSCULUS (Briq.) Bhatti & hirsuta, dendropilis et simplipilis, adaxialis glabrata. Inflorescentia Ingr., stat. nov. terminalis ad 230 mm, laxa vel densa ordinate verticillastris numerosis multifloribus; bracteae c. 4.0-5.0 x 1.0 mm, lanceolatae, hirsutae. Pogostemon section Racemosa subsection Glabriuscula Briq. Calyx tubulatus, tubo 5.0 mm longo, dentibus quinque 0.9-1.3 mm Filaments lacking moniliform hairs. A few simple hairs may be bilabiata longis, triangularibus, sub fructu incurvatis. Corolla present. purpurea, tubo ad 8.4 mm longo exserto, labio superiori lobus centralis 1.3 x 0.8 mm, labio inferiori 2.3 x 1.5 mm longo integro. 35. Pogostemon speciosus Benth. in Wall., PL asiat. rar. 1: 31 Stamina quator filamentis exsertis, pari inferiori longiori, prope (1830). antherae medianum pilis longis barbati, prope inferior villosus; Figs 9b, 25b. uniloculares. Discus non lobatus. Nuculae maturae quatuor, semi- c. Leaves 65 quaternari-spheroideus, spinifer. Stem solid, terete; hairs 8-celled, 1750 um long. ovate, x 62 mm, base cordate, apex acute, margin double crenate; hairs 6- TYPE. Philippines, Luzon, Ilocos, Norte Prov., Mt. Darna, Febru- celled, c. 1000 um long. Petiole 60 mm long; hairs 8-celled, c. 1600 ary-March 1953, Edano 18046 (BM!-holotype). (am long. Inflorescence a single terminal spike, c. 100 mm long, c. 1.0 Stem solid, terete; hairs 6-celled, c. 780 urn long. Leaves ovate, 25- dense; hairs 9-celled, c. 1700 um long; bracts small, filiform, x 4.5 5 65 x 20-35 mm, base cuneate, apex acuminate, margin crenate; x 0.2 mm, hairy. Calyx tubular, somewhat 2-lipped, 6.4 mm, veins hairs simple-uniseriate on the adaxial surface, 6-celled, c. 870 urn main veins plus a variable number of parallel minor reaching tufts of hairs within long; dendromorphic hairs and simple hairs on the abaxial surface. teeth sinuses; outside with a few scattered hairs, c. 0.9 Petiole 10-25 mm long; hairs 6-celled, 870 urn long. Inflorescence at the junction of the teeth; teeth ciliate, c. 2.3-2.6 mm long, outer hairs 5- a single terminal spike c. 230 mm long, verticils arranged regularly mm wide at base, sometimes two narrower than others; c. 2.8 x and relatively laxly; hairs 7-celled, c. 1400 urn long; bracts lanceo- celled, c. 470 um long. Corolla up to 7 mm long; lower lip 1.6 x 0.8 late, c. 4.0-5.0 x 1.0 mm, hairy. Calyx tubular, 5.0 x 4.4 mm, 1.5 mm; upper lip c. 2.5 mm across; central lobe mm. 2.7 10-veined; sparsely hairy outside, teeth and tube glabrous within; Filaments inserted at different heights, the lowest at a height of c. teeth ciliate, c. 0.9-1.3 mm long, c. 0.8-0.9 mm wide at base; outer mm in the tube; filaments c. 1 1 . 1-1 1 .6 mm long, exserted portion two hairs 6-celled, c. 1000 urn long. Corolla up to 8.4 mm long; lower lip 7.3 mm; filaments lacking moniliform hairs in the middle,

c. 1 .3 x c. 14 lobes c. 0.8 c. 2.3 x 1.5 mm; upper lip c. 3 mm across; central lobe 0.8 tomentose at the base. Style mm long; stigma mm x mm. Filaments inserted at different heights, the lowest at a height of long; ovary glandular. Disc c. 0.8 mm long. Nutlets 4, c. 800 700 3.7 mm in the tube; filaments c. 8.9-9.3 mm long, exserted portion um, ovoid, with cup-like glands. SYSTEMATICS OF POGOSTEMON 103

Fig. 9 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Allopogostemon section Racemosus: subsection Racemosus (a) P. paludosus; subsection Glabrimculus: (b) P. speciosus, (c) P. atropurpureus, (d) P. travancoricus, (e) P. reflexus; section Zygocalyx: (f) P. wightii. Scale bar 5 mm. 104 G.R. BHATTI AND M. INGROUILLE

DISTRIBUTION. India (Nilgiri (Kunda Hills), Ootacamund, on the crenate to double crenate; hairs 7-celled, 600 um long. Petiole c. 30 border of Kerala with Tamil Nadu). mm long; hairs 5-celled, c. 650 um long. Inflorescence a single terminal spike, c. 80 mm long, dense; hairs 6-celled, c. 750 um long. SPECIMENS EXAMINED. Bourne 5314 (K), Callaro s.n. (K), Clarke Calyx campanulate, 5.0 x 4.0 mm, 10-veined; a broken annulus of 10651 (K), Gamble 11611 (K), Gamble 1 1630 (K), Gamble 18389 hairs in the throat, a tuft of hairs at the sinuses; teeth ciliate, c. 1 .5- (K), Hohenacker 1225 (BM, K), Townsend & Ramamoorthy 47 (K), 1.7 mm long, 0.6-0.9 mm wide at base; outer hairs 7-celled, c. 750 Schmidt s.n. (BM), Vine 215 (BM), Wight 2028 (K). um long. Corolla up to 9.4 mm long; lower lipc. 2.3 x 1 .5 mm; upper The calyx of Pogostemon paludosus approaches that found in spe- lip c. 3 mm across; central lobe 1.1 x 1.1 mm. Filaments inserted at cies of section Zygocalyx. different heights, the lowest at a height of 3.2 mm in the tube; filaments c. 12-13 mm long, exserted portion c. 6.8 mm; filaments the c. 36. Pogostemon travancoricus Bedd., Icon. pi. Ind. or. 1: 34 glabrous towards middle, two villous towards the base. Style (1868-1874). 15.5 mm long; stigma lobes 0.8 mm. Ovary glandular. Disc c. 1 mm Figs 9d, 26a. long. Nutlets 4, c. 1200 x 1 100 um, oblong, with cup-like glands.

Stem solid, angular, glabrous. Leaves ovate, 50 x 38 mm, base DISTRIBUTION. India (Kerala, Nilgiri, Kunda Hills, Anaimalai cuneate, apex acuminate, margin crenate, lamina glabrous. Petiole c. Hills). 38 mm long, glabrous. Inflorescence a single terminal spike, c. 45 SPECIMENS EXAMINED. Beddome s.n. (BM, K), Hb. Miers s.n. mm long, lax, glabrous. Calyx tubular, 6.3 x 5.0 mm, 10-veined; (BM), Wight 2127 (K-holotype). glabrous within and outside; teeth not ciliate, c. 1.3-1.8 mm long, c. 0.9-1.1 mm wide at base. Corolla up to 9 mm long; lower lip c. 2.1 lib. Section ZYGOCALYX Bhatti & Ingr., sect. nov. 1.2 x 2 mm; upper lip c. 3.8 mm across; central lobe 1.6 x mm. Filaments inserted at different heights, the lowest at a height of 3.5 Calyx plusminusve bilabiatus, dentibus subulatibus, fimbriatus mm in the tube; filaments c. 12.8-13.9 mm long, exserted portion c. setaceus, fauci annularis pilibus. 8.4 mm; filaments glabrous towards the middle, equally villous at Calyx irregular, somewhat or markedly two-lipped, teeth awl-shaped the base. Style c. 14.1 mm long; stigma lobes c. 1.0-1.1 mm. Ovary and fringed with bristles, throat ringed by hairs, either densely or glandular. Disc c. 1 . 1 mm long. Nutlets 4, c. 1 000 x 800 jam, oblong, sparsely. with cup-like glands.

DISTRIBUTION. Southern India (Travancore). 39. Pogostemon fraternus Miq., Fl. Ned. Ind., Eerste bijv. 2: 635 (1859). SPECIMENS EXAMINED. Beddome 109 (BM, K-isotypes),/tao 3223 Figs lOa, 26c. (K). Herb up to 1 m tall; stem solid, terete, glabrous. Leaves ovate, 89 x 37. Pogostemon reflexus Benth. in A. DC., Prodr. 12: 155 (1848). 51 mm, base cuneate, apex acute, margin double dentate; hairs 5- Figs 9e, 26b. celled; c. 1000 um long. Petiole c. 45 mm long, glabrous. Inflorescence a terminal spike, c. 80 mm long, dense, with more than Stem solid, terete; hairs 6-celled, c. 610 urn long. Leaves ovate, 63 x two lateral spikes; hairs 4-celled, c. 750 um long. Calyx infundibu- 40 mm, base truncate, apex acute, margin double crenate; hairs 5- lar, c. 6.5 x 3.5 mm, 10-veined; sparsely hairy outside and with celled, c. 500 urn long. Petiole 2 1 mm long; hairs 5-celled, c. 670 urn spherical glandular trichomes equally or more common, an annulus long. Inflorescence a single terminal spike, c. 1 1 5 mm long, dense; of hairs in the throat; teeth margin, c. 2.2-3.3 mm long, c. 0.8-1.1 hairs 6-celled, c. 760 urn long. Calyx tubular, 6.8 x 6.0 mm, 10- mm wide at base; outer hairs 3-celled, c. 660 |im long. Corolla up to veined; hairy outside, an annulus of hairs in the throat; teeth ciliate, 7.5 mm long, purple- to pinkish white; lower lipc. 2.7 x 1 mm; upper c. 1.9-2.1 mm long, c. 1.0- 1.3 mm wide at base; outer hairs 5-celled, lip c. 1.5 mm across; central lobe c. 1.5 x 0.6 mm. Stamens violet; c. 560 u,m long. Corolla up to 7.7 mm long; lower lipc. 2.3 x 1 .2 mm; filaments inserted at different heights, the lowest at a height of 3.5 upper lip c. 2.6 mm across; central lobe 1.5 x 1 mm. Filaments all mm in the tube; filaments c. 8.0-8.8 mm long, exserted portion c. 4.8 inserted at a height of 2.8 mm in the tube, c. 8-10 mm long, exserted mm; filaments more or less glabrous towards the base. Style c. 1 1 portion c. 2 mm; filaments glabrous towards the middle, two villous mm long, violet; stigma lobes 0.6-0.9 mm. Disc c. 0.8 mm long. towards the base. Style c. 12 mm long; stigma lobes c. 1.1 mm. Nutlets 4, c. 1 100 x 900 um, orbicular, reticulate-foveate. Ovary glandular. Disc c. 0.6 mm long. Nutlets 4, c. 1 100 x 800 um, oblong, with cup-like glands. DISTRIBUTION. Widespread from northern India (Sikkim, Gangtok, Darjiling, Naga Hills, gorge of the Trin river, south of Saramati) to DISTRIBUTION. Sri Lanka (Kandy, Adam's Peak, Nuwara-Eliya). China (Yunnan, Szemao Forest) and Thailand (Chiang Mai, Kao SPECIMENS EXAMINED. Cramer 3483 (K), Cramer 3488 (K), Lwta Kao Kem, Korat, Kao Soi Dao-Chantatum, Doi Sutip, Kao Cramer 4028 (K), Cramer 4448 (K), Davidse & Sumithraarachchi Sunj, Tranj), Burma (Kachin, Keenam, Tennaserium, Chin Hills), 8653 (K), Moon 199 (BM), Thwaites 154 (BM), Walker s.n. (K- and Java. In dry forests, on precipitous cliffs and open ground. syntype). SPECIMENS EXAMINED. Bular s.n. (K), Clarke 21239A (K), Henry 1 1699 (K), Hooker 25 (K), Horsfield 28 (K), Hosseus 198 (BM, K), 38. Pogostemon atropurpureus Benth. in A. DC., Prodr. 12: 154 Kerr 1646 (BM, K), Kerr 3554 (BM), Kerr 4685 (BM), Kerr 9658 (1848). (BM), Kerr 15252 (BM), Kerr 16762 (BM), Kingdon-Ward 21740 Figs 9c, 25c. (BM), Lister 20 (K), Meebold 7378 (K), Moorely s.n. (K), Tun Aung Pogostemon imberbe C.H. Wright ex Hook.f., Fl. Brit. India 4: 637 & Tha Hla 3626 (K). (1885). Called 'Prao Lam' in Thailand. Miquel (1859) described the fila- Stem solid, terete; hairs 5-celled, 750 um long. Leaves elliptic to ments as glabrous. Hooker (1885) clearly mentioned that Miquel elliptic-oblong, 70 x 37 mm, base rounded, apex acute, margin found glabrous filaments in Javan plants, but noted that they were SYSTEMATICS OF POGOSTEMON 105

Fig. 10 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Allopogostemon section Zygocalyx: (a) P.fraternus, (b) P.

menthoides, (c) P. macgregorii, (d) P. litigiosus, (e) P. hirsutus, (f) P. brachystachyus, (g) P. nigrescens, (h) P. strigosus. Scale bar 5 mm. G.R. BHATTI AND M. INGROUILLE 106

Indian he examined. SPECIMENS EXAMINED. J. & M.S. Clemens 29042 (BM), J. & M.S. hairy towards the base in the plants Mukerjee towards the but Clemens 29727 J. & M.S. Clemens 32575 (BM), J. & M.S. (1940) also reported stamens were only hairy base, (BM), Clemens 40267 Press (1982) scored the filaments as hairy towards the middle. (BM). of P. (1969) regarded Pogostemon fraternus as a synonym have sometimes Keng This is a very distinct species, although specimens menthoides. He commented that the description of P. fraternus made been incorrectly identified as Pogostemon menthoides. on Horsfield's collection from G. Praoe, Java, is identical with the

off! menthoides at K. Hooker ( 1 885) treated Horsfield type specimen 42. Pogostemon macgregorii W.W. Sm. in Rec. hot. Surv. India 6: there is 28 as P. in the Flora of British India. In fact, fraternus 39(1914). another Horsfield 29, which has been considered as the specimen, Figs lOc, 27b. type of P. menthoides. Stem solid, erect; hairs 6-celled, c. 720 um long. Leaves ovate, 80 x Ned. Ind. 3: 825 25 base rounded, acute, serrate; hairs 5-celled, c. 40. Pogostemon menthoides Blume, Bijdr. fl. mm, apex margin 520 Petiole 10 mm hairs 5-celled, c. 560 um long. (1826). um long. long; Inflorescence a terminal c. 60 mm long, with 1 2 closely Figs lOb, 27a. single spike, from arranged verticils, the lowermost whorl of flowers separated Leaves 45 Stem solid, angular; hairs 5-celled, c. 600 urn long. ovate, x the rest; hairs 7-celled, c. 550 (am long. Calyx tubular-inflated, 4.0 x 23 base cuneate, acute, margin dentate; hairs 6-celled, c. mm, apex 3.6 mm, 10-veined; an annulus of hairs inside the tube, outside a few c. 670 urn 625 urn long. Petiole 6 mm long; hairs 5-celled, long. widest glandular tipped hairs present; teeth ciliate, c. 1-2 mm long, Inflorescence a terminal spike, c. 40 mm long, dense through- single tooth c. 1.0-1.3 mm wide at base; outer hairs 4-celled, c. 340 um 3.5 x 3.3 out; hairs 5-celled, c. 500 urn long. Calyx tubular, mm, x long. Corolla up to 6.0 mm long; lower lip 1.8 1.3 mm; upper lip 10-veined; outside, an annulus of hairs in the throat; teeth hairy c. 2.6 mm across; central lobe c. 1.1x0.6 mm. Filaments all inserted at outer hairs ciliate, c. 1 .2-1 .4 mm long, c. 0.5-0.8 mm wide base; at a height of 2.6 mm in the tube, c. 3.8-4.5 mm long, exserted Corolla to 4.6 mm lower c. 2.0 6-celled, c. 700 urn long. up long; lip towards the base. c. portion c. 1 . 1 mm; filaments equally hairy Style c. 1 .7 central lobe c. 1 . 1 x 0.6 mm. x 0.9 mm; upper lip mm across; c. 0.9 5.4 mm long; stigma lobes c. 0.7 mm long. Disc mm long. the lowest at a of 2 mm Filaments inserted at different heights, height reticulate-foveate. Nutlets 4, c. 1 . 1 x 1 . 1 mm, orbicular, dark brown, in the tube; filaments c. 3.5^.3 mm long, exserted portion c. 1.7 Northern Thailand Mai, mm; filaments more or less glabrous at the base. Style c. 4.5 mm DISTRIBUTION. (Pa-Ngem Chiang Nutlets c. 1 000 long; stigma lobes c. 0.5 mm. Disc c. 0.5 mm long. 4, Doichong). x 800 orbicular, reticulate-foveate. jam, SPECIMENS EXAMINED. Gam>438 (BM, K), Hansen & Smitinand Funkuoka 9659 Murata DISTRIBUTION. Widely distributed from India (Manipur, Ukhrul) 12661 (K), Iwatsuki, & Chintayungkum (K), to Indonesia andVietnam (Khu-Tu-Tri-Tay-Bac, near Chapa). Grow- 15941 (K). ing in patches at roadsides in the shade of the climax forest. 43. Pogostemon hirsutus Benth., Labial, gen. spec.: 155 (1833). ETHNOBOTANY. In Java the fragrant leaves of Pogostemon Figs lOe, 28a. menthoides are kept among clothes as an insect repellent. Leaves Stem erect, solid, angular; hairs 4-celled, c. 500 um long. SPECIMENS EXAMINED. Horsfield 29 (K-isotypes), Kingdon-Ward hairs ovate, c. 26 x 15 mm, base cuneate, apex acute, margin serrate; Petelot 51 10 18036 (BM), Kingdon-Ward 20423 (BM), (BM). c. 4-celled, c. 620 um long. Petiole c. 7 mm long; hairs 4-celled, 500 a terminal to 32 mm The filaments have been described as glabrous in various floras, but um long. Inflorescence single spike, up long, c. 5 x 4 a few moniliform hairs have been observed. This is a strongly dense; hairs 6-celled, c. 1000 um long. Calyx tubular, mm,

it with scattered hairs outside, glabrous within; scented plant with branches rooting so that forms a carpet. 10-veined; very long c. 0.7 teeth unequal, ciliate, longest tooth c. 1 .5 mm long, shortest hairs c. 820 um mm long, c. 0.5-0.7 mm wide at base; outer 5-celled, 41. litigiosus Doan in Humbert, Fl. gen. Indo-China Pogostemon 1 x 1 .2 long. Corolla up to 6.3 mm long; lower lip c. 2. mm; upper lip 4:972(1936). x 0.8 c. 2.5 mm across, hairy outside; central lobe c. 1.5 mm. Figs lOd, 27c. Filaments inserted at different heights, the lowest at a height of 2.5 c. 2.5 Stem solid, terete, rooting at the nodes; hairs 8-celled, c. 1600 urn mm in the tube; filaments c. 3.2-6.3 mm, exserted portion mm; villous at the base. c. 7.0 mm long. Leaves ovate, 54 x 34 urn, base truncate, apex acute, margin filaments all more or less equally Style Nutlets c. 450 double dentate; hairs 7-celled, c. 1 1 00 urn long. Petiole 25 mm long; long; stigma lobes c. 0.6 mm. Disc c. 0.5 mm long. 4, hairs 7-celled, 1400 urn long. Inflorescence a single terminal spike, x 300 um, oblong, reticulate-foveate. c. 60 mm long, lax throughout; hairs 7-celled, c. 1300 urn long. DISTRIBUTION. Sri Lanka (Hakgala, Nuwara-Eliya, Matale). Calyx bilabiate, 5.5 x 5.5 mm, 10-veined; with scattered long hairs outside, an annulus of hairs in the throat; teeth unequal, ciliate, c. SPECIMENS EXAMINED. Cramer 4872 (K), Macrae 446 (K- Townsend 73/ 0.8-2.3 mm long, c. 1.1-1.5 mm wide at base; outer hairs 5-celled, holotype), Simpson 9049 (BM), Thwaites 283 (BM), c. x c. 680 (am long. Corolla up to 6.5 mm long, white; lower lip 1.8 155 (K). 0.6-1.5 mm. 1.5 mm; upper lip c. 1.5 mm across; central lobe 44. Labial, 156 (1833). Filaments inserted at different heights, the lowest at a height of 2.5 Pogostemon wightii Benth., gen. spec.: 28b. mm in the tube; filaments c. 5.2-5.4 mm long, exserted portion c. 1.4 Figs 9f, mm; filaments glabrous towards the base. Style c. 7.7 mm long; Stem solid, angular; hairs 9-celled, c. 1700 um long. Leaves ovate, lobes c. 0.7-0.8 mm. Disc c. 0.5 mm long. Nutlets 4, c. 1200 stigma 50 x 43 mm, base truncate, apex acute, margin double crenate; hairs x 1000 um, orbicular, reticulate-foveate. 1400 7-celled, c. 1500 um long. Petiole 26 mm long; hairs 9-celled, Inflorescence a terminal c. 70 mm lax; DISTRIBUTION. Only found in Sabah on the slopes of Mt Kinabalu um long. single spike, long, hairs c. 1700 um tubular, 5.7 x 5.7 mm, 10- above 1000 m. Along wet forest trails and in other wet places. 8-celled, long. Calyx SYSTEMATICS OF POGOSTEMON 107 veined; sparsely hairy outside, an annulus of hairs in the throat; teeth lobe 0.5-0.9 mm wide at base. Filaments inserted at different ciliate, c. 1.3-1.5 mm long, c. 0.8-1.1 mm wide at base; outer hairs heights, the lowest at a height of 1 .7 mm in the tube; filamentsc. 3.2- 7-celled, c. 1500 jam long. Corolla up to 6 mm long; lower lip c. 2.2 3.5 mm long, exserted portion c. 1.3 mm; filaments equally villous x 1.5 mm; upper lip c. 2.5 mm across; central lobe 1.1 x 0.7 mm. towards the base. Style c. 3.9 mm long; stigma lobes c. 0.6 mm. Disc Filaments inserted at different heights, the lowest at a height of 2.4 c. 0.5 mm long. Nutlets 4, c. 800 x 600 um, obovoid, reticulate. mm in the tube; filaments c. 6.0-6.5 mm long, exserted portion c. 5.0 DISTRIBUTION. Western China (Yunnan). mm; two filaments villous at the base. Style c. 7.0 mm long; stigma lobes c. 0.4-0.5 mm. Disc c. 0.8 mm long. Nutlets 4, c. 600 x 500 SPECIMENS EXAMINED. Forrest 18590 (BM, K), Henry 9082 & urn, orbicular, reticulate-foveate with secondary reticulation. 9082T (K), Henry 1 1 174 (K-holotype), Henry 12563 (K).

DISTRIBUTION. Southern India (Nilaghiri, Kunda Hills, Malabar, 47. Pogostemon strigosus (Benth.) Benth. in A. DC., Prodr. 12: Concan). 155(1848). SPECIMENS EXAMINED. Gamble 16929 (K), Gamble 17853 (BM, Figs lOh, 29b. K), Klackenberg & Lundin 116 (K), Stocks & Law s.n. (BM), Dysophylla strigosa Benth. in Wall., PL asiat. rar. 1: 30 (1830). Schmidt 74 (BM), Wight s.n. (K). Stem solid, angular; hairs 4-celled, c. 875-960 um long. Leaves 45. Pogostemon brachystachyus Benth. in A. DC., Prodr. 12: 156 lanceolate, 41-51 x 12-20 mm, base truncate to rounded, apex (1848). acute, margin serrate; hairs 5-celled, c. 1200 um long. Petiole c. 4 Figs lOf, 28c. mm long; hairs 5-celled, c. 800-1000 um long. Inflorescence a single terminal spike, c. 70-108 mm long; hairs 4-5 celled, c. 1000 Stem solid, rooting at nodes, angular; hairs up to 6-celled, c. 900 (am um long. Calyx tubular, 4.3-5.0 x 3.2^4.2 mm, 10-veined; with long. Leaves ovate, 33 x 20 mm, base cuneate, apex acuminate, scattered hairs outside, a tuft of hairs at the sinuses within; teeth margin dentate; hairs 7-celled, c. 550 (am long. Petiole c. 6 mm long; ciliate slightly inside the margin, c. 0.9-1.7 mm long, 0.5-0.9 mm hairs 5-celled, c. 400 urn long. Inflorescence a single terminal spike, wide at base; outer hairs 3-celled, 620-750 um long. Corolla up to c. 140 mm long; hairs 6-celled, c. 630 urn long. Calyx tubular, c. 4 x 5.8 mm long; lower lip c. 2.0 x 1 . 1 mm; upper lip c. 2. 1 mm across; 3.8 mm, 10-veined; outside with a few hairs, some glandular, an central lobe 1.1 x 0.7 mm. Filaments all inserted at a height of 2.4- annulus of hairs in the throat; teeth ciliate, c. 1.3-1.6 mm long, 0.4- 2.9 mm in the tube, c. 5.0-6.6 mm long, exserted portion c. 2.3-3.5 0.6 mm wide at base; outer hairs 4-celled, c. 470 urn long. Corolla up mm; filaments all equally villous at base. Style c. 7.0-7.8 mm long; to 4 mm long, bright purple; lower lip c. 1 .5 x 1 mm; upper lip c. 1 .3 stigma lobes c. 0.9 mm. Disc c. 0.5-0.6 mm long. Nutlets 4, c. 800- mm across; central lobe 1 . 1 x 0.5 mm. Filaments inserted at different 900 x 700-800 jam, obovoid, reticulate-foveate with secondary heights, the lowest at a height of 1 .6 mm in the tube; filaments c. 3.5- reticulation or spinulose. 4.0 mm long, exserted portion c. 1 .6 mm; filaments glabrous towards northeastern the base. Style c. 5.2 mm long; stigma lobes c. 0.9 mm. Disc c. 0.5 DISTRIBUTION. India (Khasia Hills, Assam) and Bang- in and at an elevation of mm long. Nutlets 4, c. 800 x 700 urn, orbicular, reticulate. ladesh. Found roadside woods pastures 1500-1800 m. DISTRIBUTION. Northern India (Khasia, Shillong) to Burma (Sumpra Prum). A weed of open roadsides, forming mat-like colo- SPECIMENS EXAMINED. Clarke 550 1 (BM), Clarke 45456B (BM), 18763 nies with ascending verticillasters. Griffith 209 (BM), Hooker s.n. (K), Kingdon-Ward (BM), Lemann 209 (K), Wallich 1549 (BM, K-isotypes). SPECIMENS EXAMINED. Clarke 15559A (BM, K), Clarke 40303

(BM), Griffith 3967 (K-isotypes), Kingdon-Ward 20519 (BM, K). III. Subgenus DYSOPHYLLUS (Blume) Bhatti & Ingr., In Kew herbarium there are two specimens of Griffith 3967 from comb, et stat. nov. East Bengal, dated 14 November 1834, and a third specimen dated Dysophylla Blume ( 1 826), pro parte. 24 September 1850. Bentham (1832-1836) described the filaments of Pogostemon Stems often weak. Leaves sessile to subsessile, rarely petiolate, in than two at each node, linear to brachystachyus as glabrous or nearly so (filamentis nudis vel leviter whorls of more two, rarely opposite or to ovate or orbicu- barbatis). Hooker (1885), Mukerjee (1940), and all other workers linear-lanceolate elliptic elliptic-oblong, rarely fila- lar. Inflorescence dense. infundibular, < 3 mm reported this character in the same fashion. Nevertheless, the Calyx small, usually thin and exterior with ments of P. brachystachyus are clearly densely hairy towards the in length, 5-veined, translucent, usually hairy than middle. glands obvious, circumference of the flowering calyces greater > of the total length. Width of lower corolla lip usually upper, length Nutlets lower lip equal to its width at the base. mostly oblong, rarely 46. Pogostemon nigrescens Dunn in Notes R. hot. Gdn Edinb. 8: ellipsoid to ellipsoid-oblong or ovoid. 159(1913). Figs lOg, 29a. Ilia. Section DYSOPHYLLUS 30 Stem solid, angular; hairs 4-celled, c. 370 um long. Leaves ovate, Dysophylla section Oppositifoliae Benth. (1832), pro parte, x 20 mm, base cuneate, apex obtuse, margin crenate; hairs 4-celled, Dysophylla section Eudysophylla Kudo (1927), pro parte. c. 275 um long. Petiole 4 mm long; hairs 5-celled, c. 250 um long. lax dense various but Inflorescence a terminal spike, c. 170 mm long, below, Leaves two, opposite at each node, shape commonly or above, with two lateral spikes; hairs 5-celled, c. 350 um long. Calyx broadly elliptic, ovate or lanceolate, deeply pinnatifid. of campanulate, 4.0 x 4.1 mm, 10-veined; hairy outside, an annulus 48. auricularius (L.) Hassk. in Hoeven & de Vriese, hairs in the throat; teeth ciliate, c. 0.9-1.2 mm long, 0.8-0.9 mm Pogostemon Natuurl. Gesch. 10: 127 (1843). wide at base; outer hairs 4-celled, 270 (am long. Corolla up to 3.9 mm Tijdschr. Physiol. c. 1.5 across; central 11 a, 29c. long; lower lip c. 1.3 x 0.7 mm; upper lip mm Figs 108 G.R. BHATTI AND M. 1NGROUILLE

m

Fig. 11 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Dysophyllus section Dysophyllus: (a) P. auricularius, (b) P.

glabratus, (c) P. barbatus, (d) P. amaranthoides, (e) P. myosuroides, (f) P. salicifolius, (g) P. quadrifolius, (h) P. tisserantii, (i) P. mutamba, (j) P. micangensis. Scale bar 5 mm. SYSTEMATICS OF POGOSTEMON 109

Mentha auricularia Mant. 81 L., pi: (1767). 50. Pogostemon barbatus Bhatti & Ingr., sp. nov. auricularia Dysophylla (L.) Blume, Bijdr. fl. Ned. Ind. 3: 825 Figs 1 Ic, 30b. (1826). Herbae hirsutae. Caules ad nodo contractes, fumosi. Foliae oppositae, Stem procumbent and rooting at the nodes, solid, angular, contracted lamina ad 80 x 19 mm, oblonga, crenata, cuneata, hirsuta in at of the the base internodes, pinkish; hairs up to 8-celled,c. 1200 urn superficiebus ambabus folii, pilibus c. 1.25 mm, auricoloribus in to long. Leaves opposite pairs, ovate ovate-lanceolate, up to c. 50 glandulibus depressibus in lamina, petioli c. 20 mm. Inflorescentia x 30 mm, base cuneate, apex acute, margin serrate to double serrate, ad 90 mm, gracilis, dens, ad 30 verticillastros; bracteae c. 2.5 x 0.5 lamina glandular-punctate on both surfaces; hairs 6-celled, c. 2000 mm, oblanceolatae, hirsutae ad marginem. Calyx campanulatus, Petiole to urn long. up 5 mm long, densely hairy; hairs 8-celled, c. tubo 15 x 22 mm, dentibus quinque 0.4-0.5 mm longis, ciliatus, sub 2000 urn long. Inflorescence a single terminal spike, c. 80 mm long, fructu incurvatis. Corolla bilabiata purpurea, tubo ad 3.1 mm, labio dense, the flowers fragrant; hairs on stalk 7-celled, c. 1 600 urn long. superiori lobus centralis 1 .3 x 0.8 mm, labio inferiori 1 .0 x 1 .0 mm. Floral leaves narrowly elliptic. Calyx campanulate-infundibuliform, Stamina quator filamentis exserti, pari superiori affixus ad 2 mm c. 2.5 x 3.5 mm, 5-veined; glabrous within, glandular and with a few corollae tubo, pari inferiori ad 1 .5 mm, prope medianum pilis longis hairs outside; teeth ciliate, 0.5-0.7 mm long, c. 0.8-1.0 mm wide at barbati, prope inferior glabratus; antherae uniloculares. Discus non the base, incurved in fruit; outer hairs 4-celled, c. 500 urn long. lobatus. Nuculae 400 x 300 um maturae quatuor, oblongae, Corolla c. 3.5 mm long, purplish; lower lip c. 1 x 0.8 mm; upper lip reticulatae-foveatae. c. 2.8 mm across, central lobec. 0.8 x 0.6 mm. Filaments all inserted TYPE. Hong Kong, Fau Tan Valley, 12 October 1972, Hu 12319 at a height of 1.6 mm in the tube, c. 4.3-4.1 mm long, exserted (K!-holotype). portion c. 2.8 mm; filaments glabrous towards the base. Style c. 5 mm long; stigma lobes c. 0.8 mm. Disc c. 0.3 mm long. Nutlets 4, c. Stem procumbent, solid, 4-angled, contracted at base of internodes, 700 x 380 um, ellipsoid-fusiform, brown, reticulate. greyish brown; hairs 8-celled, c. 1250 um long. Leaves in opposite pairs, c. 80 x 19 mm, oblong-clavate, base cuneate, apex bluntly DISTRIBUTION. Throughout south Asia from Sri Lanka (Kandy acute, margin crenate to double crenate, lamina hirsute, glandular, Dist., Pussellawa, Uva Prov., Sabaragamuwa Prov.), India (Madras, the glands golden, sunken; hairs c. 5-celled, c. 750 um long. Petiole Assam, Sambalpur), Malaysia, Burma (Maymo Plateau), through c. 20 mm long; hairs c. 8-celled, c. 1210 um long. Inflorescence a China (Hainan, Yunnan, Canton, Hong Kong), and south to New single terminal spike, up to 90 mm long, slender, with c. 30 confluent Guinea. Common on marshy ground and in paddy fields. whorls; hairs 8-celled, c. 1150 um long. Floral leaves clavate- ETHNOBOTANY. The leaves ofPogostemon auricularius are crushed oblanceolate, c. 2.5 x 0.5 mm; hairs only on margin, up to 5-celled, with lime and applied as a poultice on the abdomen for various c. 750 um long. Calyx campanulate, c. 15 x 22 mm, 5-veined inside; ailments, such as diarrhoea, colic, worms, and a sore throat. Its use glabrous within, with ball-like glands outside; teeth ciliate, c. 0.4- for disturbances of the stomach in children is very common in 0.5 mm long, c. 0.3-0.5 mm wide at base, incurved in fruit. Corolla

1 1 x 1 central Malaya. c. 3. mm long; lower lip mm; upper lipc. 2 mm across; lobe c. 0.6 x 0.6 mm, sparsely hairy outside. Two filaments under SPECIMENS EXAMINED. Balcher 72 1 (K), Bourne 663 (K), Cramer central lobe inserted higher (c. 0.4 mm) than other two; filaments 4056 (K), Cramer 5178 (K), Ford 3 (K), Henry 1231 1 (K), Henry equal in length, c. 3.5 mm long, exserted portion c. 1.8 mm; s.n. (K), Hepper, Maxwell & Fernando 4556 (K), Hooker 344 (K), filaments glabrous at the base, hairs towards the middle of filament Jayasuriya 1454 (K), Lace 4246 (K), Lei 886 (K), McClure 9279 on one side, c. 750 um long. Style c. 4 mm long; stigma lobes c. 0.5 (K), Mooney 1827 (K), Parry 267 (K), Peng 6250 (K), Playfair 124 mm. Nutlets 4, c. 400 x 300 jam, oblong, brown, reticulate-foveate. (K), Wiakabu et al. 73782 (K). DISTRIBUTION. Hong Kong. SPECIMENS EXAMINED. Hu 5652 49. Pogostemon glabratus Chermsir. ex Press in Bull. Br. Mus. nat. (K). Hist. (Hot.) 10: 71 (1982). This species is closely allied to Pogostemon auricularius and P. lib, 30a. Figs glabratus but differs in having more dense and much longer hairs, a crenate to the leaf, filaments attached at Stem erect, 4-angled, contracted at base of internodes; hairs few, 3- longer petioles, margin different in the corolla tube, and oblong nutlets. celled, c. 250 um long, golden glands present. Leaves in opposite heights pairs, elliptic to ovate-elliptic, c. 70 x 30 mm, base cuneate, apex acute, margin dentate, lamina punctate; hairs few, white, c. 3-celled, 51. Pogostemon amaranthoides Benth. in A. DC., Prodr. 12: 153 Inflorescence a c. 300 urn long. Petiole c. 0.5 mm long. single (1848). terminal c. whorls spike, 90 mm long, densely arranged. Calyx Figs 1 Id, 30c. campanulate, c. 2 x 2.5 mm, 5-veined; glabrous within, glandular Herb up to 1 m high; stem solid, angular; hairs 4-celled, c. 310 um outside; teeth not ciliate, c. 0.5-0.7 mm long, c. 0.4-0.7 mm wide at Leaves in ovate, 1 15 x 55 mm, base cuneate, obtuse at long. opposite pairs, base. Corolla c. 2.5 mm long; lower lip c. 0.6 x 0.6 mm, dentate; hairs 4-celled, c. 325 urn central lobe apex acuminate, margin long. apex; upper lip c. 2 mm across, with a few hairs outside; Petiole c. 30 mm long, glabrous. Inflorescence a terminal spike, c. c. 0.5 x 0.6 mm. Filaments all inserted at a height of c. 1 mm in the hairs 4- 65 mm long, lax below, dense above, with two lateral spikes; tube, c. 2.5-3 mm long, exserted portion c. 1.5 mm, glabrous celled, c. 320 um long. Calyx campanulate, 2.8 x 3.6 mm, 5-veined; towards the base. c. 4 mm long; stigma lobes c. 0.3 mm. Disc Style within hairy on ribs outside, teeth hairy on margin and just margin; c. 0.3 mm Nutlets 4, c. 700 x 500 um, ellipsoid-fusiform with long. 3- teeth c. 0.5-0.7 mm long, 0.5-0.7 mm wide at base; outer hairs small attachment point, light brown, reticulate-foveate. celled, c. 250 um long. Corolla up to 2.6 mm long, pinkish white;

1 .4 central lobe 0.6 lower lipc. 0.8 x 0.7 mm; upper lipc. mm across; DISTRIBUTION. Thailand (Hi Dat and Rachaburi, Kanburi area). x 0.5 mm; lobes with a few hairs. Filaments inserted at different

the lowest at a of 1 mm in the tube; filaments c. 1.5- SPECIMENS EXAMINED. Kerr 10274 (BM), Put 1 32 (BM-holotype). heights, height 110 G.R. BHATTI AND M. INGROUILLE

c. 2 filaments all towards the base. c. 5.2 1 .9 mm long, exserted portion c. 3 mm. Style c. 3 mm long, with a mm; sparingly hairy Style c. c. Disc c. 3 Nutlets c. few hairs; stigma lobes c. 0.7 mm. Disc c. 0.5 mm long. Nutlets 4, mm long; stigma lobes 0.5 mm. mm long. 4, x 200 dark smooth. 600 x 500 jam, oblong, puncticulate. 700 um, oblong, very shiny brown,

DISTRIBUTION. Eastern Himalayas (Tibet (Ham Tawai valley), DISTRIBUTION. Western India (Bombay, Maharashtra, Sikkim, Khasia, Darjiling, Bhutan, Chukka Dist. Chukka Dzong Mahabaleshwar, Concan, Mahabaleh Hills). Nepal). SPECIMENS EXAMINED. Dalzell s.n. (K-holotype), Hooker s.n. SPECIMENS EXAMINED. Clarke 9520A (K), Clarke 24838B (K), (K), Liam s.n. in Herb. Hooker (K), Townsend 73/25 (K). Dawson 257 (BM), Gamble Till (K), Gamble 3850A (K), Gamble Rev. 2: 530 488388 (K), Grierson & Long 3227 (K), Griffith 2 1 2 (K), Hara et al. 54. Pogostemon quadrifolius (Benth.) Kuntze, gen. pi. 06306529 (BM, K), Hooker s.n. (BM), Hooker s.n. (K-syntype), (1891). Hooker s.n. (K), Kingdon-Ward 7410 (K), Manandhar 8348 (BM), Figs llg, 31c. Stainton, Sykes & Williams 9276 (BM). Mentha stellata Roxb., Hon. bengal: 44 (1814), nom. illegit. Dysophylla quadrifolia Benth. in Wall., PI. asiat. rar. 1: 30 (1830). 52. Pogostemon myosuroides (Benth.) Kuntze, Rev. gen. pi. 2: 530 D. velutina Benth. in Wall., PI. asiat. rar. 1: 30 (1830). (1891). D. rupestris Dalzell in Hooker's J. Bot. 3: 120 (1851). Figs lie, 3 la. Eusteralis quadrifolia (Benth.) Panigrahi in Phytologia 32: 478 Dysophylla myosuroides Benth. in Wall., PI. asiat. rar. 1: 30 (1830). (1976). D. rugosa Hook.f., Fl. Brit. India 4: 638 (1885). E. quadrifolia (Benth.) Majumdar in J. Bombay nat. Hist. Soc. 74: Pogostemon rugosus (Hook.f.) El Gazzar & L.Watson in Taxon 16: 385(1978, '1977'). 187(1967). Stem erect, solid, terete; hairs 3-celled, c. 437 um long. Leaves Stem erect, solid, terete, greyish because of hairs; hairs 4-celled, c. mostly in whorls of 4, sometimes in opposite pairs, lanceolate, c. 70 base hairs c. 550 um long. Leaves in opposite pairs, lanceolate, c. 40 x 7 mm, base x 12 mm, cuneate, apex acute, margin serrate; 4-celled, c. hairs similar to those of stem. cuneate, apex bluntly acute, margin sharply dentate, lamina thick, 750 um long. Petiole 4 mm long; each branch terminated a terminal densely hairy on both sides, veins strongly developed; hairs 3- Inflorescence much branched, by hairs c. celled, c. 500 um long. Petiole c. 3 mm long; hairs 4-celled, c. 560 spike, spike on main axis c. 120 mm long, dense; 4-celled, c. 2 x 2.7 um long. Inflorescence a terminal spike, c. 50 mm long, dense, with 750 um long. Calyx campanulate, mm, 5-veined; hairy teeth c. 0.4-0.5 0.4-0.6 lateral spikes, c. 30 mm long, dense; hairs 4-celled, c. 550 um long. outside, glabrous within; ciliate, mm long, c. 3 10 Corolla to Floral leaves linear-oblong. Calyx campanulate, c. 1 .3 x 1 .7 mm, 5- mm wide at base; outer hairs 3-celled, um long. up c. x 0.9 c. 2 veined; hairy outside, glabrous within; teeth more or less equal, 3.3 mm long; lower lip 0.9 mm; upper lip mm across; Filaments all inserted at a ciliate, longest tooth c. 0.3 mm long, shortest 0.2 mm long, widest central lobe 0.8 x 0.6 mm; lobes hairy. tooth c. 0.3 mm wide, narrowest tooth c. 0.2 mm wide; outer hairs 4- height of 1 .4 mm in the tube, c. 4.2^.5 mm long, exserted portion at the base. 4.5 celled, c. 430 um long. Corolla 1 .5 mm long; lower lip 0.5 x 0.5 mm; c. 2.6 mm; filaments equally hairy Style mm long; lobes c. 0.9-1 mm. Disc c. 0.3 Nutlets upper lipc. 1 mm across; central lobe c. 0.4 x 0.3 mm; a few hairs on stigma unequal, mm long. 3, the lobes. Filaments all inserted at a height of 0.5 mm in the tube, c. c. 500 x 400 jam, ellipsoid-oblong, dark brown, spinulose. 1-1.3 mm long, exserted portion c. 0.3 mm. Style 2.2 mm long, DISTRIBUTION. Northern India (Khasia, Assam, Sambalpur) and stigma lobes c. 0.7 mm. Discc. 0.2 mm long. Nutlets 4,c. 1000 x 700 Bangladesh (Chittagong), and also from Kerala (Malabar, Godavari um, ellipsoid to ellipsoid-oblong, dark brown, spinulose. Distr., Garo Hills) in southern India. In eastern Bangladesh it grows DISTRIBUTION. Eastern India (Madras, Kaumbollan Hill). on sandy soils.

SPECIMENS EXAMINED. Feicher 4724 (K), Gamble 20353 (K), SPECIMENS EXAMINED. Barber 7 1 1 5 (K), Bourne 3396 (K), Clarke 15965 Wallich 1547 (BM, K-syntypes), Wight 2130 (K), Wight 2533 (K), 19782 (K), Gamble 6739A (K), Gamble 15964 (K), Gamble Wight 2534 (K). (K), Griffiths 205 (K), Hooker 1538 (K), Hooker 1539 (K), Hooker 1540 (K), Monney 2557 (K), Parry 937 (K), Parry 1040 (K), Raza salicifolius ex El Gazzar L. 53. Pogostemon (Dalzell Hook.f.) & 786 (K), Roxburgh s.n. (K-syntype). Watson in Taxon 16: 187 (1967). Tahir et al. (1995) report a reticulate nutlet surface and a calyx only Figs llf, 31b. hairy on the teeth in material from Bangladesh. Dysophylla salicifolia Dalzell ex Hook.f., Fl. Brit. India 4: 638 (1885). 55. Pogostemon micangensis G. Taylor in/ Bot., Lond. 69 (Suppl. 2): 166(1931). Stem erect, solid, terete; hairs appressed, 2-celled, the cells more or Figs llj, 32c. less equal, c. 425 um long. Leaves in opposite pairs, linear to linear- lanceolate, c. 35 x 16 mm, base cuneate, apex cuneate, margin Aquatic herb with spreading caespitose rhizome; stem weak, terete, shallowly dentate; hairs 2-celled, c. 560 um long, basal cell slightly red-purple; hairs 4-celled, c. 325 um long. Leaves in opposite pairs, hairs 3- longer than upper cell. Petiole c. 0.5 mm long; hairs 3-celled, c. 440 ovate, 22 x 9 mm, base cuneate, apex acute, margin serrate;

1 hairs c. 200 um long. Inflorescence a single terminal spike, c. 60 mm long, celled, c. 200 um long. Petiole mm long; 3-celled, um dense; hairs 2-celled, c. 680 um long. Calyx campanulate, c. 2.5 x long. Inflorescence a single terminal spike, c. 44 mm long, dense; x 5.5 2.7 mm, 5-veined; hairy outside, especially near the base, teeth hairy hairs 2-celled, c. 430 um long. Calyx campanulate, 3.8 mm, within; teeth ciliate, c. 0.4-0.5 mm long; outer hairs 3-celled, c. 500 10-veined; glabrous or with a few hairs outside, teeth and upper part 0.8-1.4 um long. Corolla c. 3 mm long; lower lipc. 1 x 1 mm; upper lipc. 1.8 of tube hairy within; teeth ciliate, c. 0.4-1.2 mm long, mm mm across; central lobe c. 0.8 x 0.4 mm. Filaments all inserted at a wide at base; outer hairs 5-celled, c. 580 um long. Corolla up to 4.9 height of 1.2 mm in the tube, c. 3.9-4.2 mm long, exserted portion mm long, whitish violet; lower lip c. 1.1 x 1 .4 mm; upper lip c. 1 .7 SYSTEMATICS OF POGOSTEMON 111 mm across; central lobe 0.8 x 0.5 mm, hairy. Filaments inserted at This is the only species of Pogostemon which has compound leaves. different heights, the lowest at a height of 2.3 mm in the tube; filaments c. 5.7-6.0 mm long, exserted portion c. 3.4 mm; filaments glabrous towards the base. Stylec. 5.8 mm long; stigma lobesc. 0.7- Illb. Section VERTICILLATUS (Benth.) Bhatti & Ingr., 1 mm. Disc c. 0.5 mm long. Nutlets 4, c. 400 x 300 jam, ellipsoid to comb. nov. ellipsoid-oblong, reticulate-foveate. Dysophylla Blume, pro parte ( 1 826), Dysophylla section Verticillatae DISTRIBUTION. Angola (River Micango) and Cameroon. Found in Benth. (1832), Eusteralis Raf. (1847), Dysophylla subgenus the marshes of the river Micango. Chotekia Keng, pro parte (1927).

SPECIMENS EXAMINED. Gossweiller 2545 (BM-isotype), J. & A. Leaves usually linear to linear-lanceolate (P. trinervis orbicular), Raynal 12161 (BM). sessile, truncate, rounded or cuneate), more than two leaves at each node (P. andersonii two leaves). Gossweiller 2545 has calyces which are 5, 6 & 7 toothed.

58. Pogostemon lythroides (Diels) Press in Bull. Br. Mus. not. Hist. 56. Pogostemon mutamba (Hiern) G. Taylor in J. Bot., Land. 69 (Bot.) 10: 74 (1982). (Suppl. 2): 166 (1930). Figs 12a, 33a. Figs Hi, 32b. Dysophylla lythroides Diels mNotizbl. Bot. Gart. Mus. Berl. 9: 1031 mutamba Hiern, Cat. afr. pi.: 854 (1900). Geniosporum (1926).

Amphibious perennial with a polycephalic rootstock; stem weak, Stem terete, weak, glabrous. Leaves in whorls of 4, linear, violet angular; hairs 4-celled, c. 275 jam long. Leaves in opposite pairs, above, c. 88 x 10 mm, base truncate, apex acute, margin distantly to elliptic-oblong, soft-flaccid, 90 x 35 mm, base rounded, elliptic serrate, revolute, lamina with midvein sparsely hairy abaxially, hairs 6-celled, c. 600 Petiole 5 apex acute, margin dentate; |am long. lateral veins obscure on adaxial surface but distinct on abaxial mm hairs 4-celled, 300 urn long. Inflorescence a procumbent long; surface; hairs 2-celled, c. 180 um long. Inflorescence a single to ascending terminal spike, c. 60 mm long, with more than two terminal spike c. 16 mm long, glabrous. Floral leaves claviform, lateral all dense; hairs 4-celled, c. 680 urn long. Calyx spikes, spikes linear to linear-lanceolate. Calyx campanulate, c. 28 x 35 mm; 2.5 x 3.0 mm, 5-veined; densely hairy outside, teeth campanulate, outside densely hairy throughout, teeth hairy within; teeth ciliate, c. within; teeth ciliate, c. 0.5-0.7 mm long, c. 0.5-0.6 mm wide hairy 0.9-1.1 mm long, c. 0.5-0.7 mm wide at base; outer hairs 3-celled, at base; outer hairs 5-celled, c. 430 urn long. Corolla up to 5.6 mm c. 560 um long. Corolla c. 4 mm long; lower lip c. 1 .4 x 1 .4 mm; whitish lower c. 1.9 x 1.6 c. 2 mm long, violet; lip mm; upper lip c. 0.7 x 0.5 and upper lip c. 2.8 mm across; central lobe mm; upper central lobe c. 1.1 x 0.5 mm. Filaments all inserted at across, hairy; at of 1 .5 lower lip hairy outside. Filaments all inserted a height mm a of 2.8 mm in the tube, c. 5.0-5.7 mm, exserted portion c. 2.9 height in the tube, c. 4.0-4.5 mm long, exserted portion c. 2.2 mm; filaments towards the base. 7 mm long; stigma mm; glabrous Stylec. filaments glabrous at the base. Style c. 5.5 mm long; stigma lobes lobesc. 0.7 mm, violet-purple. Discc. 0.5 mm long. Nutlets 4,c. 400 0.6-0.8 mm long. Disc c. 0.4 mm long. Nutlets 4, c. 400 x 300 um, x 300 urn, reticulate-foveate. oblong, oblong, reticulate-foveate. DISTRIBUTION. Ganda and Caconda, Planalto, Angola (between DISTRIBUTION. Widespread throughout south Asia: India Benguela). (Sambalpur, Khasia), Sri Lanka, Malaysia, Sulawesi, Papua New China ETHNOBOTANY. Pogostemon mutamba is a source of starchy ed- Guinea, (Yunnan). ible tubers. SPECIMENS EXAMINED. Cramer 3996 (K), Cramer 4902 (K), 5084 Duthie 9698 Fan & Li 563 SPECIMENS EXAMINED. Gossweiler 9668 (BM), Welwitsch 5496 Cramer (K), (K), (BM), Flenley 2104 1544 (K), Hartley 9689 (K), Mooney 2139 (K), (BM-syntype), Welwitsch 5590 (BM, K-syntypes). (K), Griffith Mooney 3226 (K), Steenis 1260 (K). nov. 57. Pogostemon tisserantii (Pellegr.) Bhatti & Ingr., comb. ciliatus Bhatti & nov. Figs llh, 32a. 59. Pogostemon Ingr., sp. Figs 12b, 33b. r. Avanc. Sci. 49: 387 Dysophylla tisserantii Pellegr. in C. Ass. fr. solides bicellularibus c. (1926). Herba villosa. Caules quadrangulati, pilibus 750 um: in quoque nodus 4 folia lineares, verticillatae, ad 40 x 3 Stem weak; hairs thin, whitish, 2-celled, c. 200 urn long. Leaves in mm; cuneati et acutifolia, margine integre, pilibus bicellularibus, c. whorls of 4, deeply pinnatifid, c. 30 mm long; hairs sparse, 1 -celled, 560 um long. Inflorescentia terminalis ad c. 70 mm longa, pilibus in c. 75 um Inflorescence spikes c. 70 urn long, flowers long. bicellularibus c. 750 um long. Calyx campanulatus, c. 1 .8 x 2.5 mm; whorls. Calyx campanulate, 2 x 2.5 mm; hairy outside interrupted dentibusc. 0.4-0.5 mm, partibus interioribus pilibus, margine ciliato; teeth teeth ciliate, c. 0.6-0.7 and distinctly glandular, hairy within; c. 750 um. Corolla in partibus exterioribus villosibus, bicellularibus, c. 0.4-0.5 wide at outer hairs 4-celled, c. 250 urn mm long, mm base; 1 .5 c. 2.5 mm; labio inferiori 1 x 1 mm; labio superiori mm lato; c. 0.8 x 0.8 c. long. Corolla c. 2.2 mm long; lower lip mm; upper lip labio et inferiori ciliati. lobis superiori 0.6 x 0.6 mm; superiori 1.7 mm across, outside; central lobec. 0.5 x 0.5 mm. Filaments 3.0- hairy Stamina quator filamentis affixus ad 0.9 mm in tubo; filaments in c. 3. 1 exserted all inserted at a height of 0.6 mm the tube, mm long, c. 3.4 3.3 mm; partibus exsertibus c. 1.7 mm. Styli mm; stigmatibus c. 1.5 filaments at the base. Style c. 3.3 mm portion mm; glabrous lobibus c. 0.6 mm, hirsutibus. Disc non lobatus, 0.4 mm. Nuculae 4, lobes c. 0.7 mm. Disc c. 0.3 mm long. Nutlets 4, c. 570 long; stigma c. 600 x 400 um, oblongae, granulatae, valde flexuosa. x 400 um, ovoid, somewhat cuneate at the base, reticulate-foveate. TYPE. New Guinea (expeditie 1954-55), Netherland, New Guinea DISTRIBUTION. Chad. 2 November and Kebar valley, c. 100 km west of Manokwari, 1954, 3967 SPECIMENS EXAMINED. Jacques 1807 (K). Royen (K!-holotype). 112 G.R. BHATTI AND M. INGROUILLE

Fig. 12 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Dysophyllus section Dysophyllus: (a) P. lythroides, (b) P.

ciliatus, (c) P. linearis, (d) P. pentagonus, (e) P. andersonii, (f) P. cruciatus, (g) P. sampsonii, (h) P. yatabeanus, (i) P.faurei, (j) P. peguanus, (k) P.

koehneanus, (1) P. crassicaulis. Scale bar 5 mm. SYSTEMATICS OF POGOSTEMON 113

Stem hairs c. solid, angular, villous; 2-celled, 750 jam long. Leaves linear, c. 20 x 2 mm, base truncate, apex bluntly acute to obtuse, in whorls of 40 x 3 base 4, linear, mm, cuneate, apex acute, margin margin entire, revolute, lamina glabrous. Inflorescence a single hairs c. 560 urn Inflorescence a entire; 2-celled, long. single terminal terminal spike, c. 30 mm long, with glandular and eglandular hairs; c. 70 mm hairs 2-celled. c. 750 spike, long; (am long. Calyx hairs 3-celled, c. 250 um long. Calyx campanulate, c. 1 .2 x 2 mm;

c. 1 .8 x 2.5 campanulate, mm; hairy outside, teeth hairy within; teeth sparsely hairy outside, glabrous within; teeth ciliate, c. 0.3-0.5 mm c. 0.4-0.5 0.4-0.5 wide at the ciliate, mm long, mm base; outer hairs long, c. 0.40.5 mm wide at base, incurved in fruit, closing the c. 750 Corolla c. 2.5 villous, 2-celled, jam long. mm long; lower lip mouth of the tube; outer hairs 3-celled, c. 250 um long, basal cell

1 x 1 1 .5 central lobe x mm; upper lip mm across; 0.6 0.6 mm; upper with dark contents, glands conspicuous. Corolla c. 2.5 mm long; and lower ciliate. Filaments all inserted at a of in lip height 0.9 mm upper lip c. 1.5 mm across, lower lip c. 0.7 x 1 mm; central lobe c. the 3.0-3.3 exserted c. 1.7 tube, mm long, portion mm; filaments 0.7 x 0.9 mm wide at base; upper and lower lip with a ring of hairs towards the base. c. equally hairy Style 3.4 mm long; stigma lobes c. outside. Filaments all inserted at a height of 0.8 mm height in the Disc 0.4 c. 0.6 mm, hairy. mm long. Nutlets 4, 600 x 400 um, oblong, tube, c. 3.5-4 mm long, exserted portion c. 2.2 mm; filaments with brown, granulate 'zig-zag' channels. glabrous at the base. Style 3.5 mm long, bulbous at base; stigma lobes unequal, c. 0.7 and 1 mm. Nutlets 4, c. 700 x 400 um, oblong, DISTRIBUTION. New Guinea. Growing in meadows among tall 4-angled, light brown. grasses; fairly common. DISTRIBUTION. India (Chota Nagar, Sambalpur, Bihar, and Orissa) Pogostemon ciliatus is distinct in having hairy/ciliate stigma lobes, and Thailand (Dai Thai). the upper and lower lips of the corolla ciliate, and a nutlet surface with zigzag channels (not spinulose). SPECIMENS EXAMINED. Clarke 20438 (BM, K-isotypes), Garrett 56 (BM, K), Kerr 1465 (BM, K), Mooney 1854 (K).

60. Pogostemon linearis (Benth.) Kuntze, Rev. gen. pi. 2: 529 (1891). 62. Pogostemon andersonii (Prain) Press in Bull. Br. Mus. nat. Figs 12c, 33c. Hist. (Bot.)10:71 (1982). Figs 12e, 34b. Dysophylla linearis Benth. in A. DC., Prodr. 12: 157 (1848). Eusteralis linearis (Benth.) Panigrahi in Phytologia 32: 476 (1976). Dysophylla andersonii Prain in J. Asiat. Soc. Beng. 59: 298 (1891). E. linearis (Benth.) Majumdar in /. Bombay nat. Hist. Soc. 74: 385 Eusteralis andersonii (Prain) Majumdar in/ Bombay nat. Hist. Soc. (1978,' 1977'). 74:386(1978, '1977').

Stem erect or procumbent and rooting at the nodes, weak, reddish Stem erect, terete, villous; hairs 3-celled with longer apical cell, c. brown, glabrous or with a few hairs towards the inflorescence. 350 um long. Leaves in opposite pairs or in whorls of 3 or more, Leaves in whorls of 4, linear to linear-lanceolate, c. 50 x 5 mm, base sessile, lanceolate, c. 35 x 8 mm, base cuneate to truncate, apex acute, apex bluntly acute, margin distantly dentate, lamina sparsely acute, margin distantly dentate, lamina villous; hairs up to 3-celled hairy, the abaxial surface with sunken glands; hairs 2-celled, c. 120 with long apical cell, swollen at junction between cells, c. 440 um um long. Inflorescence a single terminal spike, c. 60 mm long; hairs long. Inflorescence a single terminal spike c. 15 mm long, dense; 2-celled, c. 250 um long. Floral bracts oblong, brownish towards the hairs 3-celled, c. 350 um long. Floral leaves linear to linear-lanceo- c. 1.8 x 2.5 villous tip. Calyx infundibular, c. 2 x 2.8 mm; densely hairy outside, late. Calyx campanulate, mm; 5-veined; outside, teeth less with especially at the teeth tips, with conspicuous brown glands, glabrous teeth with a few hairs within; more or oblong bluntly within; teeth ciliate, c. 0.5-0.9 mm long, c. 0.4-0.6 mm wide at base; acute apex, ciliate, c. 0.6-0.8 mm long, 0.4-0.6 mm wide at base; outer hairs 4-celled, c. 430 um long. Corolla c. 4.2 mm long; lower outer hairs villous, 4-celled, c. 370 um long. Corolla c. 2.4 mm long;

1 . 1 0.5 x 0.5 c. 1.1 central lobe c. 0.5 lip c. 1 .3 x 1.3 mm; upper lip c. 2.5 mm across; central lobe c. x lower lip mm; upper lip mm across; all at x and lower outside. Filaments all inserted 0.7 mm; upper and lower lip hairy outside. Filaments inserted 0.4 mm; upper lip hairy in c. 1.5-1.8 exserted a height of 1 .5 mm in the tube, c. 3^ mm long, exserted portion c. at a height of 1.2 mm the tube, mm long, at the base. c. 2.5 1.3 mm; two filaments hairy. Style c. 5.5 mm long; stigma lobes c. portion c. 0.8 mm; filaments glabrous Style mm c. c. 0.2 Nutlets c. 360 0.8-1.0 mm. Discc. 0.6 mm long, shallowly lobed. Mature nutlets 1, long; stigma lobes 0.3 mm. Disc mm long. 4, reticulate-foveate. c. 1000 x 800 um, ellipsoid to ellipsoid-oblong, tan, reticulate- x 230 um, oblong, dark brown, foveate. DISTRIBUTION. Northeastern India (Sikkim). DISTRIBUTION. Western China (Dirang-Dzong, Yunnan), Bhutan. SPECIMENS EXAMINED. Anderson s.n. (K-holotype). SPECIMENS EXAMINED. Bodinier s.n. (K), Clarke 45731 (BM), 63. cruciatus (Benth.) Kuntze, Rev. gen. pi. 2: 529 Forrest 11747 (BM, K), Forrest 16059 (K), Forrest 25146 (K), Pogostemon (1891). Henry 12628 (K), Kingdon-Ward 14260 (BM). Figs 12f, 34c. Rev. 61 . Pogostemon pentagonus (C.B. Clarke ex Hook.f.) Kuntze, Dysophylla cruciata Benth. in Wall., PI. asiat. rar. 1: 30 (1830). gen.pl. 2:529(1891). Eusteralis cruciata (Benth.) Panigrahi in Phytologia 32: 478 (1976), Figs 12d, 34a. non Dvsophvlla cruciata sensu Hemsl. ex F.B. Forbes & Hemsl. non D. cruciata sensu Dunn Dysophylla pentagona C.B. Clarke ex Hook.f., Fl. Brit. India 4: 641 (1890), (1915). in nat. Hist. Soc. 74: 385 '(1885). E. cruciata (Benth.) Majumdar J. Bombay Eusteralis pentagona (C.B. Clarke ex Hook.f.) Panigrahi in (1978/1977'). 32: 477 (1976). Phytologia Stem erect, solid, terete, hirsute; hairs 5-celled, c. 1200 um long. E. (C.B. Clarke ex Hook.f.) in J. Bombay nat. pentagona Majumdar Leaves in whorls of 4-5, linear, c. 30 x 4 mm, base cuneate, apex Hist. Soc. 74: 386 (1978, '1977'). obtuse to bluntly acute, margin entire, revolute; hairs 4-celled, c. 1000 um Inflorescence a terminal c. 100 mm Stem procumbent, solid, terete, glabrous. Leaves in whorls of 4, long. single spike, 114 G.R. BHATTI AND M. INGROUILLE long; hairs stalked, 5-celled, c. 1200 mm long. Calyx campanulate, 4 mm long; lower lip c. 1.2 x 1.2 mm; upper lip c. 2 mm across; c. 1.8x2.7 mm; tube and teeth very hairy outside, teeth hairy within; central lobe c. 0.9 x 0.7 mm; upper and lower lip hairy outside. teeth ciliate, c. 0.5-0.7 mm long, c. 0.4-0.6 mm wide at base; outer Filaments all inserted at a height of 1.5 mm in the tube, c. 4.4-5.0 hairs 5-celled, c. 750 mm long. Corolla up to c. 2.7 mm long; lower mm long, exserted portion c. 2.5 mm; filaments all glabrous at the lip c. 1 x 1 mm; upper lip c. 1.8 mm across; central lobe c. 0.8 x 0.5 base. Style c. 5.5 mm long; stigma lobes 0.8-1 mm long. Disc c. 0.4 mm; upper and lower lip hairy outside. Filaments all inserted at a mm long. Nutlets 4, c. 900 x 600 um, oblong, light brown, reticulate- height of c. 1 . 1 mm in the tube, c. 3.6-4.6 mm long, exserted portion foveate with small depression at the base. c. 3 mm; filaments hairy towards the base. Style c. 4 mm long; DISTRIBUTION. Japan (Honshu, Shikoku, Kyushu) and Korea. stigma lobes c. 0.7 mm. Disc c. 0.2 mm long. Nutlets 4, c. 600 x 400 urn, oblong, basal part protuberant, light brown, smooth. SPECIMEN EXAMINED. Midutoranowo s.n. (K-isotype).

DISTRIBUTION. Widespread from eastern India and the Himalayas 66. Pogostemon faurei (H. Lev.) Press in Bull. Br. Mus. nat. Hist. (Orissa, Sambalpur, Nepal, Assam, Chota Nagpur, Nagaland, (Bot.) 10: 73 (1982). Manipur), China (Yunnan) to Thailand. Figs 12i, 35c. SPECIMENS EXAMINED. Clarke 1 8059 (BM), Clarke 343 1 2A (K), Dysophylla faurei H. Lev. in Reprium Spec. nov. Regni veg. 9: 248 Forrest 25140 (K), Hensen, Seidenfaden & Smitinand 10824 (K), (1911). Hensen, Seidenfaden & Smitinand 1 1008 (K), Meebold 6002 (K), 1855 4234 & Meeboldl6l2(K), Mooney (K), Mooney (K), Strachey Stem weak, glabrous. Leaves in whorls of 4, linear, c. 24 x 3 mm, Winterbottom 1541 (K-isotypes). base cuneate, apex acute, margin dentate, lamina glabrous. Inflores- cence a single terminal spike, c. 25 mm long; hairs 3-celled, c. 250 64. Press in Bull. Br. Mus. nat. Pogostemon sampsonii (Hance) |im long. Calyx campanulate, 2 x 2.5 mm; glabrous or sparsely hairy Hist. (Hot.) 10:74(1982). outside with very fine hairs, teeth hairy within; teeth ciliate, c. 0.5 35a. Figs 12g, mm long, widest tooth c. 0.8 mm wide at base, narrowest tooth c. 0.4 mm wide at base; outer hairs 1-celled, c. 100 um long. Corolla up to Dysophylla sampsonii Hance in Annls Sci. nat. 5: 284 (1866). 3.3 mm long; lower lip 1 x 1 mm; upper lip c. 2 mm across; central Eusteralis sampsonii (Hance) Panigrahi mPhytologia 32: 478 ( 1 976). lobe c. 0.6 x 0.5 mm. Filaments all inserted at a height of c. 1.4 mm Stem hairs c. 250 um solid, terete, sparsely hairy; 3-celled, long, in the tube, c. 3.5-3.8 mm long, exserted portion c. 1.9 mm; violet. Leaves in whorls of c. 20 x 5 base 3, linear-lanceolate, mm, filaments glabrous at the base. Style c. 4.5 mm long; stigma lobes c. obtuse to lamina truncate, apex bluntly acute, margin serrate, gla- 1 mm long. Disc c. 0.3 mm long. Nutlets 4, c. 410 x 305 um, oblong, brous. Inflorescence a terminal c. 20 single spike, mm long, dense, reticulate-punctate. stalk glabrous. Floral leaves elliptic-ovate. Calyx campanulate, c. DISTRIBUTION. Korea. In rice paddies. 1 .5 x 2.4 mm; tube and teeth hairy outside; teeth ciliate, c. 0.4-0.5 c. 0.5-0.6 mm wide at outer hairs c. 1 80 mm long, base; 2-celled, jam SPECIMENS EXAMINED. Esquiz 199 (E), Faurie 760 (E-holotype), Corolla c. 1 .7 mm lower c. 0.6 x 0.6 c. long. long; lip mm; upper lip Taquet l\n (E). 1 .6 mm across; central lobe 0.5 x 0.5 mm; upper and lower lip hairy outside. Filaments all inserted at a height of 0.8 mm in the tube, inner 67. Press in Bull. Br. Mus. nat. Hist. pair c. 3.3 mm long, outer pair 2.8 mm long, exserted portion c. 2.4 Pogostemon peguanus (Prain) 10: 74 mm; filaments glabrous at the base. Style c. 3.5 mm long; stigma (Bot.) (1982). 36a. lobes c. 0.4-0.5 mm. Disc 0.2 mm long. Nutlets 4, c. 640 x 450 um, Figs 12j, oblong, attachment point protruded, reticulate-foveate. Dysophylla peguana Prain in J. Asiat. Soc. Beng. 59: 298 (1891). DISTRIBUTION China. Stem erect, 4-angled, weak; hairs appressed, 1-celled, c. 370 um Leaves in whorls of c. 60 x 3 base ETHNOBOTANY. Pogostemon sampsonii has been used to poison long. 4, linear, mm, truncate, apex acute to lamina with flies in China (Von Reis, 1973). bluntly obtuse, margin entire, revolute, appressed hairs; hairs 1-celled, c. 400 um long. Inflorescence a single terminal SPECIMENS EXAMINED. Hance 10946 (BM, K-isotypes). spike, c. 80 mm long; stalk with glandular and eglandular hairs; eglandular hairs 1 -celled, c. 400 um long, glandular hairs 3-celled, c. 65. Pogostemon yatabeanus (Makino) Press in Bull. Br. Mus. nat. 250 um long. Calyx campanulate, c. 1.8 x 2.4 mm; hairy outside Hist. (Bot.)10:74(1982). with glandular and eglandular hairs, teeth hairy within, the hairs Figs 12h, 35b. unicellular; teeth ciliate, c. 0.4-0.6 mm long, 0.4-0.6 mm wide at base; outer eglandular hairs 1-celled, c. 400 um long, glandular hairs Dysophylla yatabeana Makino in Bot. Mag., Tokyo 1: 55 (1898). 3-celled, c. 250 jam long. Corolla c. 2.5 mm long; lower lip 1 x 0.8 Eusteralis yatabeana (Makino) Panigrahi in Phytologia 32: 478 mm; upper lip c. 1.5 mm across; central lobe 0.7 x 0.5 mm; upper (1976). and lower lip hairy outside. Flaments all inserted at a height of 1 mm E. yatabeana (Makino) Murata in Acta phytotax. geobot. Kyoto 33: in the tube, c. 3.5-3.8 mm long, exserted portion c. 2.3 mm; filament 370(1982). glabrous at the base. Style 4.5 mm long; stigma lobes c. 0.8-0.9 mm. Stem solid, terete, glabrous. Leaves in whorls of 3, linear, c. 80 x 5 Disc c. 1 mm long. Nutlets 4, c. 500 x 300 um, oblong with mm, base truncate, apex acute, margin distantly serrate, lamina protuberant base, light brown, smooth. sparsely hairy and with golden glands; hairs 2-celled, c. 150 um DISTRIBUTION. Southeastern Asia in Burma (Pegu, Lahamauge) long. Inflorescence a single terminal spike, c. 50 mm long, stalk and Thailand (Pak Zong Chai). glabrous. Floral leaves linear. Calyx campanulate, c. 2.5 mm long; outside with many hairs and glands, teeth hairy within; teeth ciliate, SPECIMENS EXAMINED. Kunz 2401 (K-syntype), Kunz 2405 (K- c. 0.5-0.7 mm long; outer hairs 3-celled, c. 370 um long. Corolla c. syntype), Lace 2899 (K), Collector unknown 8123 (K). OF SYSTEMATICS POGOSTEMON 115

68. Pogostemon koehneanus (Muschl.) Press in Bull. Br. Mus. not. Pogostemon benthamianum (Hance) Kuntze, Rev. gen. pi. 2: 530 Hist. (Bot.)10:74(1982). (1891). Figs 12k, 36b. Aquatic herb; stem angular, weak, few hairs at nodes and towards the koehneana in Dysophylla Muschl. Reprium nov. Spec. Regni veg. 4: inflorescence; hairs 3-celled, c. 280 um long. Leaves in whorls of up 269(1907). to 6, linear, 50 x 2 mm, base truncate, apex acute, margin dentate,

hairs 1 c. lamina with Stem weak; -celled, 430 um long. Leaves in whorls of 3, glabrous, golden glands sunken in the abaxial surface, c. 22 x adaxial surface covered with lanceolate, 4 mm, base cuneate, apex acute, margin serrate; a white crystalline powder. Inflores- hairs 1 c. 310 cence a terminal the axis -celled, |am long. Inflorescence a single terminal spike, spike, 60-160 mm long, main sometimes

1 shorter than the c. 50 mm long; hairs -celled, c. 430 urn long. Calyx campanulate, branches, with more than two lateral spikes; hairs on stalk c. 1 .5 x 2.3 mm; hairy outside, teeth hairy within; teeth ciliate, c. 0.5 few, 3-celled, c. 287 um long. Calyx infundibular, c. 1 .2 x 1 .7 x 0.5 incurved in outside with stout mm, fruit; outer hairs 2-celled, c. 370 urn long, mm; densely hairy throughout relatively hairs and

basal cell obvious. c. 1 round also teeth c. 0.4 x 0.4 swollen, glands Corolla .7 mm long; lower lip glands present, glabrous within; ciliate, outer hairs c. 0.7 x 0.7 mm; upper lip c. 1 .3 mm across; central lobe 0.5 x 0.4 mm; mm, spreading; 2-celled, 200 um long, basal cell Corolla to lips sparsely hairy. Filaments all inserted at a height of 0.7 mm in the glandular. up 1.5 mm long; lower lip 0.5 x 0.6 mm, few

hairs 1 tube, c. 2.4-2.7 mm long, exserted portion c. 1.4 mm; filaments outside; upper lipc. mm across, ciliate at margin, few hairs on outer central lobe x glabrous towards the base. Style c. 2.4 mm long; stigma lobes c. 0.5 surface; 0.5 0.5 mm. Filaments all inserted at a of 0.6 in c. 1 mm. Disc c. 0.2 mm long. Nutlets 4, c. 500 x 300 urn, oblong with height mm the tube, .5 mm long, exserted portion c. 0.6 filaments pointed base, brown, shiny, smooth-lineate. mm; equally hairy towards the base. Style c. 2 mm long; stigma lobes 0.4-0.5 mm. Disc c. 0.2 mm long. Nutlets 4, c. 548 x DISTRIBUTION. Thailand. 419 um, ellipsoid to ellipsoid-oblong, light brown, granular. SPECIMENS EXAMINED. Clarke 25 (K), Hosseus 704 (BM-isotype). DISTRIBUTION. Bangladesh (Sylhet, Chutia Nagpur), India, and China (Canton). In pools. According to Bentham (1848: 157) it is also found in southeast the 69. Pogostemon crassicaulis (Benth.) Press in Bull. Br. Mus. nat. Asia, Philippines, and Australia. Hist. (Bot.) 10:73(1982). ETHNOBOTANY. Vernacular name 'Pansinga baha'. The roots are Figs 121, 36c. used for the female complaint 'Sitka'. Grows on the edges of Dysophylla crassicaulis Benth. in Wall., PL asiat. rar. 1: 30 (1830). terraces. Eusteralis crassicaulis (Benth.) Panigrahi in Phytologia 32: 476 SPECIMENS EXAMINED. Sampson 443 (E), Sampson s.n. (K), (1976). Sinclair 3831 (E), Watt9S6\ (E), 24-8-1887. s.n. (K), Wallich 1543 Annual herb; stem up to 40 cm, erect, solid, terete, rooting at the (K-syntype), Wallich 1544 (BM, K-syntypes). nodes, glabrous. Leaves in whorls of 3-5, linear, c. 30 x 6 mm, base cuneate, apex acute, margin dentate, lamina brick red, sparsely 7 1 . Pogostemon pumilus (Graham) Press in Bull. Br. Mus. nat. Hist. hairy; hairs 2-celled, c. 250 urn long. Inflorescence branches termi- (Bot.) 10: 74 (1982). nating in a single spike, spike on main axis c. 80 mm long, lateral Figs 13b, 37b. spikes up to 40 mm long, stalk with glandular and eglandular hairs; hairs to c. 300 urn c. up 3-celled, long. Calyx campanulate, 1.5x2 Mentha pumila Graham in Edinb. New phil. J. 4: 393 (1828). mm; hairy outside with glandular and eglandular hairs, teeth hairy Dysophylla pumila (Graham) Benth. in Wall., PL asiat. rar. 1: 30 within; teeth ciliate, c. 0.5-0.7 mm long, c. 0.4-0.6 mm wide; outer (1830). eglandular hairs 2-4 celled, c. 250 jam long, basal cell of glandular Eusteralis pumila (Graham) Panigrahi in Phytologia 32: 477 (1976). hairs x swollen. Corollac. 2.2 mm long, violet; lower lip 0.5 0.5 mm; E. pumila (Graham) Majumdar in J. Bombay nat. Hist. Soc. 74: 385 upper lip c. 1.2 mm across; central lobe c. 0.5 x 0.3 mm; upper lip (1978, '1977'). hairy outside. Filaments all inserted at a height of c. 1 mm in the tube, c. 2.0-2.4 mm long, exserted portion c. 1.2 mm; filaments Stem erect, terete, weak, glabrous. Leaves in whorls of 4, linear to glabrous at base. Style c. 3 mm long; stigma lobes c. 0.4 mm. Disc linear-lanceolate, c. 20 x 3 mm, base truncate, apex acute, margin c. 0.2 mm long. Nutlets 4, c. 800 x 300 jam, oblong, basal part serrate, lamina glabrous. Inflorescence a single terminal spike up to protuberant, brown, shiny, the surface more or less undulate. 15 mm long, glabrous; floral leaves clavate, c. 1.7 x 0.7 mm; bracts spathulate, up to 1 mm long, hairy at margin. Calyx campanulate, c. DISTRIBUTION. Bangladesh (Sylhet), India (Allahabad, Baharaich, 1.6 x 2.5 mm; glabrous within, hairy outside, mainly on the teeth; Motipur), and southeastern Asia to Vietnam. Rooting in the black teeth ciliate, c. 0.4-0.5 mm long, c. 0.4-0.5 mm wide at base; outer mud of swamps in roadside ditches. hairs 2-3-celled, c. 200 ^im long, some glandular with dark basal x c. SPECIMENS EXAMINED. Couder s.n. (K), Panigrahi 6588 (E), cell. Corolla c. 0.8 mm long; lower lip c. 0.5 0.5 mm; upper lip c. x 0.5 outside. Squires 249 (K), Wallich 1545 (BM, K-isotypes). 1 .2 mm across; central lobe 0.4 mm; upper lip hairy Filaments all inserted at a height of 0.9 mm in the tube, c. 1 .7-2 mm long, exserted portion c. 1.1 mm; filaments glabrous towards the 70. verticillatus (Roxb.) Bhatti & Ingr., comb. nov. Pogostemon base. Style c. 2.5 mm long; stigma lobes c. 0.2-0.4 mm. Disc c. 0.2 Figs 13a, 37a. mm long. Nutlets 4, c. 500 x 400 um; oblong with protuberant base, brown, Mentha verticillata Roxb., Hort. Bengal: 44 (1814). reticulate-punctate. verticillata Benth. in PI. asiat. rar. 1: 30 Dysophylla (Roxb.) Wall., DISTRIBUTION. Nepal to Bangladesh. (1830), pro parte. s.n. Wallich 1546 D. ramosissima Benth. in Wall., PL asiat. rar. 1: 30 (1830). SPECIMENS EXAMINED. Stone (E-isotypes), (E, D. benthamiana Hance in Ann. Sc. Nat. V, 5: 234 (1866), pro parte. K). 116 G.R. BHATTI AND M. INGROUILLE

Fig. 13 Dissected corollas, calyces and bracts, showing inner surface. Pogostemon subgenus Dysophyllus section Verticillatus: (a) P. verticillatus, (b) P.

pumilus, (c) P. trinervis, (d) P. helferi, (e) P. pressii, (f) P. stocksii, (g) P. erectus, (h) P. stellatus, (i) P. deccanensis, (j) P. aquaticus. Scale bar 5 mm.

72. Pogostemon trinervis Chermsir. ex Press in Bull. Br. Mus. nat. wide at base; outer hairs 2-celled, c. 500 |im long, glandular or Hist. (Bot.) 10: 74 (1982). eglandular. Corolla c. 1 .5 mm long; lower lip c. 0.5 x 0.5 mm; upper all Figs 13c, 37c. lip c. 1.1 mm across; central lobe 0.4 x 0.4 mm. Filaments inserted at a height of 0.7 mm in the tube, three filaments c. 1 .5-1 .7 Stem erect, thin; hairs 2-celled, c. 180 jam long, apical cell some- mm long, exserted portion c. 1 . 1 mm; filaments glabrous at the base. times hook-shaped. Leaves in whorls of 3, more or less orbicular, c. Style c. 1.7 mm long; stigma lobes c. 0.4 mm. Disc 0.3 mm long. 7x5 mm, base rounded, apex obtuse, margin dentate, lamina with Nutlets 4, c. 600 x 400 urn, oblong with protuberant base, tan, shiny, short stiff hairs on both sides; hairs 2-celled, c. 180 jam long. smooth. Inflorescence branches terminated by a single spike, spike on main axis c. 20 mm stalk with and hairs; hairs long, glandular eglandular DISTRIBUTION. Thailand (Hui-Taleng, Korat, Pak-tang-chai). 2-celled, c. 250 urn long. Calyx campanulate, 1.5 x 2 mm, very hairy outside, teeth hairy within; teeth ciliate, c. 0.5 mm long, 0.4-0.5 mm SPECIMENS EXAMINED. Alam 8115 (K), Put 2223 (BM-holotype). SYSTEMATICS OF POGOSTEMON 117

73. helferi (Hook.f.) Press in Bull. Br. Mus. nat. Hist. Pogostemon Stem solid, terete, glabrous. Leaves in whorls of 10 or more, linear, (Bot.) 10: 73 (1982). c. 16x2 mm, base truncate, apex acuminate, margin entire, lamina Figs 13d, 38a. glabrous. Inflorescence a single terminal spike, c. 45 mm long; bracts cup-shaped, with fine hairs at the and Dysophylla helferi Hook.f., Fl. Brit. India 4: 640 (1885). tip margin. Calyx campanulate, pentagonous, c. 1 x 2 mm; sparsely hairy outside, Eusteralis helferi (Hook.f.) Panigrahi in Phytologia 32: 477 (1976). glabrous within, the teeth glandular; teeth ciliate, c. 0.4 x 0.4 mm, E. helferi (Hook.f.) Majumdar in J. Bombay nat. Hist. Soc. 74: 385 incurved in fruit; outer hairs 2-celled, c. 200 um Corolla c. 1 .4 (1978, '1977'). long. mm long; lower lip 0.5 x 0.5 mm; upper lip 1.4 mm across; central Stem solid, more or less a few hairs towards 4-angled, glabrous, the lobe 0.5 x 0.5 mm, with a few hairs and many glands. Filaments all inflorescence; hairs 2-celled, c. 160 Leaves in whorls of inserted at a jam long. 4, height of 0.5 mm in the tube, c. 2.5-2.8 mm long, c. 30 x 5 base linear, mm, truncate, apex acute, margin dentate, exserted portion c. 1.9 mm; filaments glabrous towards the base. lamina Inflorescence a glabrous, punctate. terminal spike, c. 20 mm Sty lee. 2.4 mm long; stigma lobes c. 0.4 mm. Discc. 0.2 mm. Nutlet with 4-5 lateral floral leaves to c. x long, spikes; wide, triangular-ovate 1, 600 200 um, narrowly oblong, light brown, smooth. bracts or rhombic; broadly narrowly spathulate, c. 1 mm long. Calyx DISTRIBUTION. Southwestern India (Bombay, Concan). campanulate, c. 1.6x2 mm, hairy outside and with white, globose within; teeth ciliate and at the c. 0.5 glands, glabrous hairy tips, mm SPECIMENS EXAMINED. Stocks s.n. in Herb. Hooker (K-holotype). long, c. 0.3-0.4 mm wide at base; outer hairs 2-celled, c. 160 um

Corolla c. 2.5 mm lower c. 0.7 x 0.7 c. 76. long. long; lip mm; upper lip Pogostemon erectus (Dalzell) Kuntze, Rev. gen. pi. 2: 530 1 .5 mm across; central lobe c. 0.5 x 0.5 mm; lobes with a few hairs (1891).

outside. Filaments all inserted at 1 a height of mm in the tube, c. 2.5 Figs 13g, 39a. mm long, exserted portion c. 1 mm; filaments glabrous towards the Dysophylla gracilis Dalzell in Hooker's J. Bot. 2: 337 (1850), non base. Style 2.8 mm long; stigma lobes c. 0.6-0.7 mm. Discc. 0.3 mm Pogostemon gracilis Hassk. (1843). long. Nutlets 4, c. 300 x 250 um, oblong, brown, shiny, smooth. Eusteralis gracilis (Dalzell) Panigrahi in Phytologia 32: 476 (1976). DISTRIBUTION. India (Bengal) and Burma (Tenasserim). E. erecta (Kuntze) Panigrahi in Phytologia 32: 477 (1976). E. erecta (Kuntze) Majumdar in J. Bombay nat. Hist. Soc. 74: 386 SPECIMENS EXAMINED. Heifer 194 (BM, E), Heifer 3968 (K- (1978, '1977'). holotype). E. gracilis (Dalzell) Majumdar in J. Bombay nat. Hist. Soc. 74: 386 (1978, '1977'). 74. Pogostemon pressii Panigrahi in Taxon 33: (1989). E. tomentosa var. gracilis Bennet & Raizada in Indian Forester 108: Figs 13e, 38b. 303(1982).

Dysophylla griffithii Hook.f., Fl. Brit. India 4: 641 (1885). Stem erect, terete, brown to purplish, sparsely hairy; hairs 3-celled, Eusteralis 1 griffithii (Hook.f.) Panigrahi inPhytologia 32: 477 ( 976). c. 375 um long. Leaves in whorls of 10 or more, linear, c. 2.5 x 1 mm, E. griffithii (Hook.f.) Majumdar in 7. Bombay nat. Hist. Soc. 74: 386 base truncate, apex acute, margin entire, revolute; hairs few, up to 3- (1978, '1977'). celled, c. 250 um long. Inflorescence a single terminal spike, c. 70 Pogostemon griffithii (Hook.f.) Press in Bull. Br. Mus. nat. Hist. mm long; hairs 3-celled, c. 370 um long; floral leaves claviform, two (Bot.) 10: 73 (1982). hairs at apex giving appearance of a cobra head with exserted Stem erect, slender, glabrous. Leaves in whorls of 4, linear, 40 x 5 tongue; bracts filiform-spathulate, c. 1-2 mm long. Calyx mm, base rounded, apex acute, margin distantly serrate, lamina campanulate, c. 1 .7 x 2. 1 mm; very hairy outside, especially in upper glabrous, glandular. Inflorescence a single terminal spike, c. 60 mm half, teeth hairy within; teeth ciliate, c. 0.5-0.6 mm long; outer hairs long; hairs 2-celled, c. 250 urn long. Calyx obconic, c. 1 .2 x 2.5 mm; white, up to 3-celled, c. 430 pm long. Corolla up to 2.4 mm long; outside the teeth with whitish hairs above, the tube glabrous, gla- lower lip c. 0.6 x 0.6 mm; upper lip c. 1 .4 mm across; central lobe c. brous within; teeth ciliate, c. 0.4 x 0.5 mm, incurved in fruit; hairs 3 0.5 x 0.4 mm; lobes hairy. Filaments all inserted at a height of c. 1 celled, c. 370 jam long. Corolla up toe. 1.7 mm long; lower lipc. 0.5 mm in the tube, c. 2.5-2.7 mm long, exserted portion c. 1 mm; x 0.7 mm; upper lip c. 1 .7 mm across; central lobe c. 0.5 x 0.4 mm. filaments hairy from middle to connective of anthers, all equally Filaments all inserted at a height of c. 0.7 mm in the tube, c. 2 mm hairy at the base. Style c. 3.5 mm long; stigma lobes c. 0.7 mm. Disc long, exserted portion c. 1 mm; filaments glabrous at the base. Style c. 3 mm long, unequal with one arm elongated. Nutlets 4, c. 400 x c. 2.2 mm long; stigma lobes c. 0.3 mm. Discc. 0.2 mm long. Nutlets 200 um, oblong, tan, smooth. 4, c. 500 x 400 um, oblong with attachment point at base, dark DISTRIBUTION. India (Bombay). brown, smooth. SPECIMENS EXAMINED. Dalzell s.n. (K-holotype), Hooker s.n. DISTRIBUTION. India (Madras) and China. (K). SPECIMENS EXAMINED. Chattaya 5224 (K), Gamble 13748 (K), Griffiths 3968 (K-holotype). var. diplolobatus Bhatti & Ingr., var nov.

a var. erecto, tow bilobo dignoscenda. 75. Pogostemon stocks!! (Hook.f.) Press in Bull. Br. Mus. nat. Hist. (Bot.) 10: 74 (1982). TYPE. India: Kerala; Karimbam, 23 December 1 980, Ansari 69986 Figs 13f, 38c. (K!-holotype).

Dysophylla stocksii Hook.f., Fl. Brit. India 4: 642 (1885). Differs from var. erectus in having a 2 armed disc. Eusteralis stocksii (Hook.f.) Panigrahi inPhytologia 32: 478 (1976). Rev. 2: 429 E. stocksii (Hook.f.) Majumdar in J. Bombay nat. Hist. Soc. 74: 386 77. Pogostemon stellatus (Lour.) Kuntze, gen. pi. (1978, '1977'). (1891). 118 G.R. BHATTI AND M. INGROUILLE

DISTRIBUTION. India Madras). Figs 13h, 39b. (Bombay, Mysore, Madras Herb. Mentha stellata Lour, in Flora cochinch. 2: 361 (1790). SPECIMENS EXAMINED. Dalz s.n. (K-holotype),

rar. 1: 30 1 1982 1407 Dysophylla stellata (Lour.) Benth. in Wall., PL asiat. (K), Ramamoorthy (K). (1830), non Mentha stellata Roxb. (1814). Press in Bull. Br. Mus. D. benthamiana Hance in Ann. Sc. Nat. V, 5: 234 (1866), pro parte. 79. Pogostemon aquaticus (C.H. Wright) Hist. 10: 73 D. peguana auct., non Prain (1891). nat. (Bot.) (1982). Eusteralis stellata (Lour.) Panigrahi in Phytologia 32: 477 (1976). Figs 13j, 40. E. malabarica in J. Bombay nat. Hist. Soc. 74: (Lour.) Majumdar 5: 45 1 Elsholtzia aquatica C.H.Wright in Dyer, Fl. Trop. Afr. ( 1900). 385 (1978, '1977'), nom. inval. Stem terete, hollow, glabrous except at the nodes; hairs 4-celled; c. Stem solid, terete, with short internodes, puberulent; hairs 3-celled, 430 um long. Leaves in whorls of 4, linear, c. 18x6 mm, base base c. 3 10 um long. Leaves in whorls of up to 10, linear, 9x2 mm, cuneate, apex acute, margin entire, glandular, lamina more or less truncate, apex acuminate, margin entire, revolute; hairs 2-celled, c. glabrous, a few hairs on the midrib on abaxial surface. Inflorescence c. 55 125 um long. Inflorescence a single terminal spike, mm long; a terminal spike with six lateral spikes, all in a single whorl; terminal bracts linear-clavate or spathulate, c. 1 mm long. Calyx campanulate, more spike with long peduncle c. 70 mm long and a dense spike of white hairs c. 1 .5 x 2.2 mm, tube and teeth hairy within and with or less the same length, first whorl separated, lateral spikes c. 30 mm outside; teeth ciliate, c. 0.4-0.5 mm long, 0.4-0.5 mm wide at base, long, equalling peduncles. Floral leaves linear, c. 2 mm long. Calyx incurved in fruit; outer hairs 2-celled, c. 190 um long. Corolla c. 1 .7 campanulate, c. 2 x 3 mm; glabrous but glandular within, hairy 1 .5 central mm long; lower lip c. 0.6 x 0.6 mm; upper lip mm across; outside, more densely above to sparsely at the base; teeth ciliate, c. Filaments all lobe c. 0.5 x 0.5 mm; upper and lower lip hairy outside. 0.7 mm long, 0.5-0.7 mm wide at base; outer hairs 2-celled, c. 310 inserted at a of 0.8 mm in the tube, two under central lobe height 1 x 1 .2 um long. Corolla c. 3 mm long; lower lip .2 mm, strigose c. 3.3 mm two on lower side c. 3 mm long, exserted (inner) long, lobe c. 1 x 0.7 lobes within; upper lip c. 2 mm across; central mm; c. 2.4 mm; filaments puberulent at the base and with portion ciliate at margin. Filaments all attached at a height of 1 .2 mm in the moniliform hairs from middle to connective of anthers. Style c. 3.4 tube, c. 2.5-3.2 mm long, exserted portion c. 2.2 mm; filaments mm long; stigma lobes c. 0.9 mm. Disc c. 0.2 mm long, unequal. glabrous at the base. Style c. 4 mm long; stigma lobes c. 0.7 mm. Nutlets 4, c. 600 x 350 um, oblong, tan, smooth. Disc 0.4 mm long. Nutlets 4, c. 900 x 600 um, oblong, dark brown DISTRIBUTION. India (Bombay, Calicut, Karnataka, Mysore, Ma- to black, smooth. dras), Bangladesh, and Hong Kong. In rice paddies. DISTRIBUTION. East Africa, Zimbabwe, Mozambique (Unangu to SPECIMENS EXAMINED. Dalz s.n. (K), Gibbs 7725 (K), Heynes Lake Shireva), and Malawi (Dedza, Chongoni forest). 1542(K),Lore/ros.n.(BM-holotype),/?a026(K), Wight 2\3\ (K), SPECIMENS EXAMINED. Fanshawe F8512 (K), Johnson 15 (K- 2136 Wight (K). 8530 type), Salubeni 1780 (K), Salubeni 1888 (K), Wight (K). Tahir et al. (1995) report orbicular nutlets with protruberances in material from Bangladesh. ACKNOWLEDGEMENTS. Dr G.R. Bhatti was supported by a studentship from Shah Abdul Latif University, Khaipur, Pakistan. We would like to thank 78. Pogostemon deccanensis (Panigrahi) Press in Bull. Br. Mus. J.R. Press, R. Harley, A. Paton, I. Hedge, and staff at The Natural History nat. Hist. (Bot.) 10: 73 (1982). Museum and the Royal Botanic Gardens, Kew and Edinburgh. Figs 13i, 39c.

Dysophylla tomentosa Dalzell \nHooker's J. Bot. 2: 337 (1850), non tomentosa Hassk. Pogostemon (1844). REFERENCES Eusteralis deccanensis Panigrahi in Phytologia 32: 475 (1976). E. tomentosa (Dalzell) Panigrahi in Phytologia 32: 477 (1976). Bakhuizen van den Brink, R. C. & van Steenis, C.G.G.J. 1 963. A note onPogostemon E. tomentosa (Dalzell) Majumdar in J. Bombay nat. Hist. Soc. 74: Desf. and Dysophylla Blume. Taxon 17: 235-236. 385(1978/1977'). Bentham, G. 1829. Bot. Reg. 15: t. 1282 & t. 1300. of Indian Labiatae enumerated in the 1 830. Synopsis of the genera and species Stem terete, with short internodes, brownish violet; hairs 3-celled, c. Plantae Asiaticae catalogue of the collections in DrWallich's charge. In N. Wallich, 250 Leaves in whorls of 6-8 or more, linear, c. 8 x 2 mm, |im long. Rariores 1: 28-31. London. more or less hairs base truncate, apex acute, margin entire, revolute; 1832-1836. Labiatarum genera et species. London. Prodromus naturalis 3-celled, c. 437 um long. Inflorescence a single terminal spike, c. 50 1848. Labiatae. In A.P. de Candolle, systematis regni 12: 27-603. mm long, tapering towards apex; hairs 5-celled, c. 1000 um long; vegetabilis Bhatti, G.R. 1995. Systematic! o/Pogostemon Desf. and related genera. Ph.D. thesis. bracts c. 1-2 mm c. filiform-spathulate, long. Calyx campanulate, London. 1 .7 x 3 teeth and tube and outside; mm; hairy within, hairy glandular Blume, C.L. 1826. Bijdragen tot de Flora van Nederlandsch Indie 3. Batavia. at K. Die Natiirlichen teeth ciliate, the longest c. 0.6-0.7 mm long, 0.4-0.6 mm wide Briquet, J. 1 897. Labiatae. In A. Engler & Prantl, Pflanzenfamilien 183-374. base; outer hairs 4-celled, c. 500 |im long. Corolla c. 2.3 mm long; 4(3a): Leipzig. misc. Brown, N.E. 1909. Pogostemon rogersii in Diagnoses africanae: XXXII. Bull. 1 lobec. 0.5 lower lip 0.9 x 0.9 mm; upper lipc. .7 mm across; central Inf. R. hot. Gdns. Kew [1909]: 376-380. x 0.5 and lower outside. Filaments all inserted mm; upper lip hairy Desfontaines, R.L. 1815. Description d'un nouveau genre de Labiee. Mem. Mus. Hist. at a height of c. 1 mm in tube, c. 3.2-3.5 mm long, exserted portion nat. Paris!: 154-156. an from floral minutiae and c. 2.2 mm; two filaments with long moniliform hairs from the El-Gazzar,A. & Watson, L. 1967. Consequences of escape floristics in certain Labiatae. Taxon 16: 186-189. middle to connective of anthers, two filaments only hairy at middle, Hartwell, J.L. 1982. Plants used against cancer. A survey. Lawrence, Mass. all at the base. c. 3.8 lobes c. 0.5 glabrous Style mm long; stigma aut reformatae Hasskarl, J.K. 1 842. Plantarum genera et species novae javanenses. c. 0.3 mm. Discc. 0.2 mm long, uneven, with one lobe elongated mm Flora, Jena 25, Beib. 2(1-2): 1-32. in notch between nutlets. Nutlets 4, c. 500 x 450 um, orbicular, tan Hermann, P. 1717. Musaeum Zeylanicum 3. Lugduni Batavorum. 1885. Flora British India 4. London. with whitish notch at the base, smooth to reticulate. Hooker, J.D. of SYSTEMATICS OF POGOSTEMON 119

Keng, H. 1 969. Flora Malesianae precursores 48. A revision of Malesian Labiatae. 1984. Nomenclatural notes on Pogostemon Desf. Taxon 33: 102. Gdns' Bull., Singapore 24: 13-180. Prain, D. 1891. Noviciae Indicae. III. Some additional species of Labiatae. J. Asiat.

1978. Labiatae. In C.G.G.J. Steenis (Ed.) Flora Malesiana I, 8(3): 301-394. Soc. Beng.59: 294-318. Kudo, Y. 1927. Labiatarum sino-japonicarum prodromus. Mem. Fac. Sci. Agric. Press, J.R. 1982. Taxonomic studies in the Labiatae tribe Pogostemoneae. Bull. Br. Taihoku imp. Univ. 2(2): 37-332. MU.I. not. Hist (Bot.) 10: 1-89. 2. 2. Kuntze, O. 1891. Revisio generum plantarum Leipzig. Rafinesque-Schmaltz , C.S. 1837. Flora Telluriana Philadelphia. Linnaeus, C. 1747. Flora Zeylanica. Holmiae. Roxburgh, W. 1814. Hortus Bengalensis. Serampore.

1767. Mantissa plantarum 1. Holmiae. 1 832. Flora Indica 3. Serampore. Loureiro, J. 1790. Flora Cochinensis 2. Ulyssipone Ku in phi us, G.E. 1750. Herbarium Amboinense 6. Amsterdam. Merrill, E.D. 1912. Phil. Journ. Sc. Bot. 7: 347. Steam, W.T. 1992. Botanical Latin, 2nd ed. London.

Miquel, F.A.W. 1 859. Flora Indiae Batavae 2. Amsterdam. Tahir, S.S., Khanam, M. & Husain, S.Z. 1995. A micromorphological study of Mukerjee, S.K. 1940. A revision of the Labiatae of the Indian empire. Rec. hot. surv. Pogostemon Desf. species (Lamiaceae) from Bangladesh. Pakist. J. Bot. 27: 73-82. Ind. 14(1): 1-228. Von Reis, S.A. 1973. Drugs and foodfrom little known plants. Cambridge, Mass. Panigrahi, G. 1 976.Taxonomic notes on certain taxa ofAsiatic angiosperms. Phytologia Wu, C.Y. & Li, H.W. 1975. Some changes of botanical names in Chinese Labiatae. 32: 473-479. Ada phytotax. sin. 13: 72-95. 120 G.R. BHATTI AND M. INGROUILLE

Fig. 14 Nutlet and nutlet surface, (a) P. benghalensis, (b) P. villosus, (c) P. plectranthoides. SYSTEMATICS OF POGOSTEMON 121

'

Fig. 15 Nutlet and nutlet surface, (a) P. pubescens, (b) P. gardneri, (c) P. cristatus. 122 G.R. BHATTI AND M. INGROUILLE

Fig. 16 Nutlet and nutlet surface, (a) P. paniculatus, (b) P. purpurascens, (c) P. cablln. SYSTEMATICS OF POGOSTEMON 123

Fig. 17 Nutlet and nutlet surface, (a) P. nepetoides, (b) P. heyneanus, (c) P. nelsonii. 124 G.R. BHATTI AND M. INGROUILLE

Fig. 18 Nutlet and nutlet surface, (a) P. glaber, (b) P. hispidus, (c) P. wattll. SYSTEMATICS OF POGOSTEMON 125

P. P. elsholtzioides. Fig. 19 Nutlet and nutlet surface, (a) P. championii, (b) formosanus, (c) 126 G.R. BHATTI AND M. INGROUILLE

Fig. 20 Nutlet and nutlet surface, (a) P. tuberculosus, (b) P. griffithii, (c) P. dielsianus. SYSTEMATICS OF POGOSTEMON 127

P. mollis. Fig. 21 Nutlet and nutlet surface, (a) P. nilagaricus, (b) P. vestitus, (c) 128 G.R. BHATTI AND M. INGROUILLE

Fig. 22 Nutlet and nutlet surface, (a) P. mtundatus, (b) P. rupestris, (c) P. rogersii. SYSTEMATICS OF POGOSTEMON 129

P. (c) P. velatus. Fig. 23 Nutlet and nutlet surface, (a) P. petiolaris, (b) philippinensis, 130 G.R. BHATTI AND M. INGROUILLE

Fig. 24 Nutlet and nutlet surface, (a) P. williamsii, (b) P. membranaceus, (c) P. elatispicatus. SYSTEMATICS OF POGOSTEMON 131

Fig. 25 Nutlet and nutlet surface, (a) P. paludosus, (b) P. speciosus, (c) P. atropurpureus. 132 G.R. BHATTI AND M. INGROUILLE

Fig. 26 Nutlet and nutlet surface, (a) P. travancoricus, (b) P. reflexus, (c) P.fraternm. SYSTEMATICS OF POGOSTEMON 133

P. Fig. 27 Nutlet and nutlet surface, (a) P. menthoides, (b) P. macgregorii, (c) litigiosus. 134 G.R. BHATT1 AND M. INGROUILLE

Fig. 28 Nutlet and nutlet surface, (a) P. hirsutus, (b) P. wightii, (c) P. brachystachyus. SYSTEMATICS OF POGOSTEMON 135

^ '

- -' >* ^.

auricularius. Fig. 29 Nutlet and nutlet surface, (a) P. nigrescens, (b) strigosus, (c) 136 G.R. BHATTI AND M. 1NGROUILLE

Fig. 30 Nutlet and nutlet surface, (a) P. glabratus, (b) P. barhatus, (c) P. amamnlhoides. SYSTEMATICS OF POGOSTEMON 137

P. Fig. 31 Nutlet and nutlet surface, (a) P. myosumides, (b) P. salicifolius, (c) quadrifolius. 138 G.R. BHATTI AND M. INGROUILLE

Fig. 32 Nutlet and nutlet surface, (a) P. tisserantii, (b) P. mutamba, (c) P. micangensis. SYSTEMATICS OF POGOSTEMON 139

Fig. 33 Nutlet and nutlet surface, (a) P. lythroides, (b) P. ciliatus, (c) P. linearis. 140 G.R. BHATTI AND M. INGROUILLE

Fig. 34 Nutlet and nutlet surface, (a) P. pentagonus, (b) P. andersonii, (c) P. cruciatus. SYSTEMATICS OF POGOSTEMON 141

P. P. Fig. 35 Nutlet and nutlet surface, (a) P. sampsonii, (b) yatabeanus, (c) faurei. 142 G.R. BHATTI AND M. INGROUILLE

Fig. 36 Nutlet and nutlet surface, (a) P. peguanus, (b) P. koehneanus, (c) P. crassicaulis. SYSTEMATICS OF POGOSTEMON 143

Fig. 37 Nutlet and nutlet surface, (a) P. verticillatus, (b) P. pumilus, (c) P. trinervis. 144 G.R. BHATTI AND M. INGROUILLE

Fig. 38 Nutlet and nutlet surface, (a) P. helferi, (b) P. pressii, (c) P. stocksii. SYSTEMATICS OF POGOSTEMON 145

Fig. 39 Nutlet and nutlet surface, (a) P. erectus, (b) P. stellatus. (c) P. deccanensis. 146 G.R. BHATTI AND M. INGROUILLE

Fig. 40 Nutlet and nutlet surface of Pogostemon aquaticus. SYSTEMATICS OF POGOSTEMON 147 SYSTEMATIC INDEX

names are in Accepted roman and synonyms in italic; new names and principal references are in bold.

Blume 107 Dysophylla 78, tomentosa (Dalzell) Panigrahi 1 18 helferi (Hook.f.) Press 1 16, 117, 144 section Kudo 107 Eudysophylla 78, yatabeana (Makino) Murata 1 14 heyneanus Benth. 93, 94, 123 section Benth. 107 Oppositifoliae 78, yatabeana (Makino) Panigrahi 1 14 hirsutus Benth. 105, 106, 134 section Verticillatae Benth. 111 78, Geniosporum mutamba Hiern 1 1 1 hispidus Prain 94,95, 124 subgenus Chotekia Keng 78, 1 1 1 Hyssopus cristatus Lam. 78, 92 imberbe C.H. Wright ex Hook.f. 104 andersonii Prain 1 1 3 Mentha auricularia Blanco 92 intermedius Benth. 89 auricularia (L.) Blume 109 auricularia L. 78, 109 koehneanus (Muschl.) Press 112, 115, 142 benthamiana Hance 115, 118 cablin Blanco 92 linearis (Benth.) Kuntze 112, 113, 139

crassicaulis 115 1 1 Benth. 112, pumila Graham 5 litigiosus Doan 105, 106, 133 cruciata Benth. 113 secunda Roxb. 91 lythroides (Diels) Press 111, 112, 139

cruciata sensu Dunn 113 stellata Lour. 1 1 8 macgregorii W.W. Sm. 105, 106, 133 cruciata sensu Hemsl. ex F.B. Forbes & Hemsl. 1 13 stellata Roxb. 110 membranaceus Merr. 101, 102, 130

faureiU. LeY 114 verticillata Roxb. 1 1 5 menthoides Blume 105, 106, 133 gracilis Dalzell 1 17 Origanum benghalense Burm.f. 89 micangensis G. Taylor 108, 110, 138 griffithii Hook.f. 117 indicum Roth 94 mollis Benth. 98, 99, 127 helferi Hook.f. 117 Perilla polystachya D. Don 94 mutamba (Hiern) G. Taylor 108, 111, 138 koehneana Muschl. 1 15 Pogostemon Desf. 77, 86 myosuroides (Benth.) Kuntze 108, 110, 137 linearis Benth. 113 section Barbata Briq. 78 nelsonii Doan 93, 94, 123 lythroides Diels 1 1 1 section Racemosa Benth. 78, 97 nepetoides Stapf. 92, 93, 123 myosuroides Benth. 78, 110 subsection Glabriuscula Briq. 102 nigrescens Dunn 105, 107, 135

peguana auct. 118 subgenus Allopogostemon Bhatti & Ingr. 97 nilagiricus Gamble 97, 98, 127

peguana Prain 1 14 section Racemosus (Benth.) Bhatti & Ingr. 97 paludosus Benth. 102, 103, 131 pentagona C.B. Clarke ex Hook.f. 1 13 subsection Racemosus 97, 98, 101, 103 paniculatus (Willd.) Benth. 90, 92, 122 pumila (Graham) Benth. 115 subsection Glabriusculus (Briq.) Bhatti & parviflorus Benth. 89 quadrifolia Benth. 110 Ingr. 102, 103 patchouly Pellet. 92

ramosissima Benth. 1 1 5 section Zygocalyx Bhatti & Ingr. 103, 104, 105 patchouly sensu Hook.f. 94

rugosa Hook.f. 78, 110 subgenus Dysophyllus (Blume) Bhatti & Ingr. 107, peguanus (Prain) Press 1 12, 114, 142 rupestris Dalzell 110 108 pentagonus (C.B. Clarke ex Hook.f) Kuntze 112,

salicifolia Dalzell ex Hook.f. 1 10 section Dysophyllus 107, 1 1 2 113, 140

sampsonii Hance 114 section Verticillatus (Benth.) Bhatti & Ingr. 79, petiolaris Benth. 98, 99, 129 stellata (Lour.) Benth. 118 111, 116 philippinensis S. Moore 100, 101, 129

stocksii Hook.f. 117 subgenus Pogostemon sensu Bhatti & Ingr. 89, 90, plectranthoides auct. pro maj., non Desf. 89 strigosa Benth. 107 93,95 plectranthoides Desf. 90, 91, 120

tisserantii Pellegr. 1 1 1 subgenus Paniculatae Benth. 78 pressii Panigrahi 116, 117, 144

tomentosa Dalzell 1 1 8 amaranthoides Benth. 108, 109, 136 pubescens Benth. 90, 91, 121

velutina Benth. 1 10 andersonii (Prain) Press 1 12, 113, 140 pumilus (Graham) Press 115, 1 16, 143

verticillata (Roxb.) Benth. 1 15 aquaticus (C.H. Wright) Press 1 16, 118, 146 purpurascens Dalzell 92, 93, 122

yatabeana Makino 1 14 atropurpureus Benth. 103, 104, 131 purpuricaulis Dalzell 89 Elsholtzia aquatica C.H.Wright 118 auricularius (L.) Hassk. 107, 108, 135 quadrifolius (Benth.) Kuntze 108, 110, 137

cristatus (Willd.) Willd. 92 barbatus Bhatti & Ingr. 108, 109, 136 reflexus Benth. 103, 104, 132 paniculata Willd. 92 battakianus Ridl. 96 reticulatus Merr. 79

Eusteralis Raf. 78 benghalensis (Burm.f.) Kuntze 89, 90, 1 20 rogersii N.E. Br. 98, 99, 128

andersonii (Prain) Majumdar 1 1 3 benthamianum (Hance) Kuntze 1 15 rotundatus Benth. 98, 100, 128

crassicaulis (Benth.) Panigrahi 1 15 brachystachyus Benth. 105, 107, 134 rugosus (Hook.f.) El Gazzar & L. Watson 1 10 cruciata (Benth.) Majumdar 113 brevicorollus Y.Z. Sun 79 rupestris Benth. 98, 99, 128 cruciata (Benth.) Panigrahi 113 cablin (Blanco) Benth. 92, 93, 122 salicifolius (Dalzell ex Hook.f.) El Gazzar & L. deccanensis Panigrahi 1 1 8 championii Prain 95, 96, 125 Watson 108, 110, 137

1 141 erecta (Kuntze) Majumdar 1 17 ciliatus Bhatti & Ingr.lll, 12, 139 sampsonii (Hance) Press 112, 114, Benth. 131 erecta (Kuntze) Panigrahi 1 17 crassicaulis (Benth.) Press 115, 142 speciosus 102, 103,

1 145 gracilis (Dalzell) Majumdar 1 17 cristatus Hassk. 90, 91, 121 stellatus (Lour.) Kuntze 16, 117,

stocksii Press 1 144 gracilis (Dalzell) Panigrahi 1 17 cruciatus (Benth.) Kuntze 112, 113, 140 (Hook.f.) 16, 117,

1 Benth. 135 griffithii (Hook.f.) Majumdar 117 deccanensis (Panigrahi) Press 16, 118, 145 strigosus (Benth.) 105, 107, suavis Ten. griffithii (Hook.f.) Panigrahi 117 dielsianus Dunn 95, 97, 126 92, 94 szemacensis S.J. Press 79 helferi (Hook.f.) Majumdar 1 17 elatispicatus Bhatti & Ingr. 101, 102, 130 (C.Y. Wu & Hsuan) Bhatti helferi (Hook.f.) Panigrahi 1 17 elsholtzioides Benth. 95, 96, 125 tisserantii (Pellegr.) & Ingr. 108, 111, linearis (Benth.) Majumdar 113 erectus (Dalzell) Kuntze 1 16, 117, 145 138

1 18 linearis (Benth.) Panigrahi 113 var. diplolobatus Bhatti & Ingr. 117 tomentosa Hassk. Li 79 travancoricus Bedd. 132 malabarica (Lour.) Majumdar 1 18 falcatus (C.Y. Wu) C.Y. Wu & H.W. 103, 104, Chermsir. ex Press 143 pentagona (C.B. Clarke ex Hook.f.) Majumdar 1 13 faurei (H. Lev.) Press 112, 114, 141 trinervis 116, Press 79 pentagona (C.B. Clarke ex Hook.f.) Panigrahi 1 13 formosanus Oliver 95, 96, 125 tsiangii (Y.Z. Sun) 132 tuberculosus Benth. 97, 126 pumila (Graham) Majumdar 1 1 5 fraternus Miq. 104, 105, 95, Benth. 129 pumila (Graham) Panigrahi 115 frutescens Graham 89 velatus 100, 101, 121 verticillatus Bhatti & 115, 1 16, 143 quadrifolia (Benth.) Majumdar 1 10 gardneri Hook.f. 90, 91, (Roxb.) Ingr. vestitus Benth. 127 quadrifolia (Benth.) Panigrahi 1 10 glaber Benth. 93, 94, 124 98, 99, Press 136 villosus Benth. 89. 90, 120 sampsonii (Hance) Panigrahi 1 14 glabratus Chermsir. ex 108, 109, wattii C.B. Clarke 124 stellata (Lour.) Panigrahi 118 gracilis Hassk. 1 17 95, 96,

Press 1 17 Benth. 103, 106, 134 stocksii (Hook.f.) Majumdar 117 griffithii (Hook.f.) wightii 126 williamsii Elmer 100, 101, 130 stocksii (Hook.f.) Panigrahi 1 17 griffithii Prain 95,97, C.Y. Y.C. 97 Press 112, 114, 141 tomentosa (Dalzell) Majumdar 1 18 var. latifolius Wu & Huang yatabeanus (Makino)

Bulletin of The Natural History Museum Botany Series

Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address.

Volume 20 Studies in Hypericum: validation of new names. N.K.B. No. 1 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. Robson. 1993. Pp. 67-70. Brathys (part 2) and 30. Trigynobrathys. N.K.B. Robson. 1990. Generic monograph of the Asteraceae-Anthemideae. K. 12 Pp. 1-151, 22 figs, 46 maps. Facsimile edition. 49.25 Bremer and C.J. Humphries. 1993. Pp. 71-177. figs. 43.25

No. 2 The marine flora of Namibia: its distributions and algal Volume 24 affinities. G.W. Lawson, R.H. Simons and W.E. Isaac. 1990. No. 1 Pre-Linnaean references for the Macaronesian flora found in Pp. 153-168, 1 fig, 7 plates. Leonard Plukenet's works and collections. J. Francisco-Ortega, The infrageneric classification of Gentiana (Gentianaceae). T.- A. Santos-Guerra and C.E. Jarvis. Pp. 1-34. N. Ho and S.-W. Liu. 1990. Pp. 169-192, 13 figs. Studies on the lichen genus Sticta (Schreber) Ach.: II. Revision of Piper (Piperaceae) in the New World. 2. The Typification of taxa from Swartz's Prodromus of 1788. D.J. taxonomy of Piper section Churumayu. M.C. Tebbs. 1990. Pp. Galloway. Pp. 35-^8. 193-236, 49 figs. 34.10 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 4. 21 Volume Genera L-O. D.M. John, G.W. Lawson, J.H. Price, WF. No. 1 Historical and taxonomic studies in the Titanoderma genus Prud'homme van Reine and W.J. Woelkerling. Pp. 49-90. in the British Isles. Y.M. Chamber- (Rhodophyta, Corallinales) Studies on the Cretan flora 3. Additions to the flora of lain. 1991. 1-80, 247 42.35 Pp. figs. Karpathos. N.J. Turland and L. Chilton. Pp. 91-100. 43.25 collections of the Thorn cv. No. 2 Early Holy (Crataegus monogyna No. 2 Observations on the benthic marine algal flora of South A.R. 1991. 1 Biflora). Vickery. Pp. 81-83, fig. Georgia: a floristic and ecological analysis. D.M. John, P.J.A. A taxonomic of the referred to the study species ascomycete Pugh and I. Tittley. Pp. 101-114. B. 1991. 76 genus Leptorhaphis. Aguirre-Hudson. Pp. 85-192, Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical figs. species (excluding Australia). D.J. Galloway. Pp. 1 15-160. and identification of The typification Calymperes Morphology and ecology of seedlings, fruits and seeds of crassilimbatum Renauld & Cardot (Musci: Calymperaceae). Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp. L.T. Ellis. 1991. Pp. 193-194, 1 fig. 42.35 161-172. A study of Bixa (Bixaceae), with particular reference to the leaf Volume 22 undersurface indumentum as a diagnostic character. R.E. and N.C. Garwood. 173-180. 43.40 No. 1 An account of southern Australian species of Lithophyllum Dempsey Pp. (Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. Campbell. 1992. Pp. 1-107, 63 figs. 41.25 Volume 25

No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. Pp. No. 2 Palynological evidence for the generic delimitation of Sechium 1-94. (Cucurbitaceae) and its allies. J.L. Alvarado, R. Lira-Saade & William Roxburgh's St Helena plants. Q.C.B. Cronk. Pp. 95-98. J. Caballero. 1992. 109-121. Pp. 43.40 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 3. No. 2 Seaweeds of the western coast of tropical Africa and adjacent Genera H-K. J.H. Price, D.M. John & G.W. Lawson. 1992. pp. islands: a critical assessment. IV. Rhodophyta (Florideae) 5. 123-146. Genera P. G.W. Lawson, W.J. Woelkerling, J.H. Price, W.F. Two new species of Solatium section Geminata (Solanaceae) Prud'homme Van Reine and D.M. John. Pp. 99-122. 1992. from Cerro del Torr in western Colombia. S. Knapp. Pp. A new species of Odontorrhynchos (Orchidaceae, 123-125. 147-152. Spiranthinae) from Boliva. D.L. Szlachetko. Pp. Fissidens ceylonensis Dozy & Molkenb. (Musci: Linnaeus's interpretation of Prospero Alpine's De plantis L.T. Fissidentaceae) and some allied taxa from southern India. exoticis, with special emphasis on the flora of Crete. N.J. Ellis. 1992. Pp. 153-156, 2 figs. Turland. Pp. 127-159. 43.40 New species of Piper (Piperaceae) from Central America. M. Book review. M.G. Gilbert. P. 161. Tebbs. 1992. Pp. 157-158. Studies on the Cretan flora 1. Floristic notes. N.J. Turland. Volume 26 No. 1 A of Chaetoceros 1992. Pp. 159-164. morphological study species from the of the Pacific ocean of Studies on the Cretan flora 2. The Dianthus juniperinus (Bacillariophyta) plankton Mexico. D.U. Hernandez-Becerril. 1996. 1-73. 43.40 complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169. Pp. 41.25 No. 2 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. Myriandra to 28. Elodes. N.K.B. Robson. 1996. Pp. 75-217. Volume 23 43.40

No. 1 Revision of Piper (Piperaceae) in the New World 3. The taxonomy of Piper sections Lepianthes and Radula. M.C. Volume 27 Tebbs. 1993. 1-50, 18 Pp. figs. No. 1 Notes on the diatom species Tetracyclus castellum (Ehrenb.) for the and of Mounting techniques preservation analysis Grunow with a description of Tetracyclus pseudocastellum diatoms. S.J. Russell. 1993. 51-54. 1 fig. 43.25 Pp. nov. sp. D.M. Williams. Pp. 1-6. from No. 2 New taxa of Gentiana (Gentianaceae) from Western China and A new species of Calymperes (Musci: Calymperaceae) Peninsular L.T. Ellis. 7-10. the Himalayan region. T.-N. Ho and S.-W. Liu. 1993. Pp. 55- Malaysia. Pp. A of the tribe 60. 2 figs. phylogenetic conspectus Hyoscyameae New combinations, names and taxonomic notes on Gentianella (Solanaceae). A.L. Hoare and S. Knapp. Pp. 1 1-30. Pteroidea: Solanaceae. (Gentianaceae) from South America and New Zealand. T.-N. A revision of Solanum section 3 1-73. 1997. 43.40 Ho and S.-W. Liu. 1993. Pp. 61-66. S. Knapp and T. Helgason. Pp. CONTENTS

77 Systematics of Pogostemon (Labiatae) G.R. Bhatti and M. Ingrouille

jlletin of The Natural History Museur BOTANY SERIES

Vol. 27, No. 2, November 1997